The Nevrorthidae , mistaken at all times : phylogeny and review of present knowledge ( Holometabola , Neuropterida , Neuroptera )

This monographic review of the Nevrorthidae Nakahara, 1915, covers all 19 validly described, extant species worldwide that belong to one of the smallest families of the order Neuroptera. The family embraces four genera: Nevrorthus Costa, 1863 (with five species occurring in the Mediterranean region), Austroneurorthus Nakahara, 1958 (with two species restricted to eastern Australia), Nipponeurorthus Nakahara, 1958 (with 11 species from eastern Asia: Japanese islands, mainland China, Taiwan), and Sinoneurorthus Liu, H. Aspöck & U. Aspöck, 2012 (with one species recorded from mainland China). A comprehensive taxonomical treatment of all extant taxa is presented, including the scant available biological data. Distribution maps for all species are provided. A phylogenetic analysis based on morphological data from both extant and extinct taxa was performed. Austroneurorthus, together with Nevrorthus and some Eocene Baltic amber genera, form a monophylum. The disjunct distribution of modern nevrorthid genera demonstrates the relictual nature of the family and points to a historical biogeography that could have led to the formation of the present distribution pattern. Future discovery of fossil material might substantiate these claims.


Scope
Deutsche Entomologische Zeitschrift is an international peer-reviewed journal of systematic entomology. It publishes original research papers in English on systematics, taxonomy, phylogeny, comparative and functional morphology, as well as biogeography of insects. Other arthropods are only considered where of relevance to the biology of insects. The geographical scope of the journal is worldwide. Priority is given to revisional work and comprehensive studies of phylogenetic, biological or zoogeographical relevance. The journal also welcomes review articles pertaining to systematics and biology of insects.
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Introduction
The family Nevrorthidae comprises only 19 described extant species assigned to four genera -with an extremely disjunct distribution (U. Aspöck and H. Aspöck 1994, 2007, Liu et al. 2012, 2014 -and nine described fossil species assigned to five genera from the Eocene Baltic amber (Wichard 2016). In addition, there is record of an undescribed putative nevrorthid from the mid-Cretaceous Burmese amber (mentioned in Makarkin 2016, based on a photograph in Xia et al. 2015).
The eidonomically inconspicuous adults are nonetheless impressive due to their excessively shaped male genital sclerites that are of high phylogenetic relevance. The aquatic larvae are equipped with a complex joint ("Rollengelenk") between head and pronotum (Zwick 1967), and the archaic head capsule has played a key role in understanding the phylogeny of Neuroptera. The aquatic pupa (Malicky 1984) is unique among Neuroptera and Neuropterida. The phylogenetic position of Nevrorthidae is controversial (Wang et al. 2016). The aim of the present paper is to summarize the accumulated knowledge on Nevrorthidae and to hypothesize on phylogenetic relationships of the family internally and within the order Neuroptera.

Historical overview
The odyssey of Nevrorthidae from nowhere to a phylogenetic key position in the context of landmarks in neuropterology (U. Aspöck and H. Aspöck 2010b) follows a unique pathway -though "mistaken at all times" -as addressed in the title. Mucropalpus fallax Rambur, 1842, the first described nevrorthid, was originally placed in Hemerobiidae (Rambur 1842). Costa (1863) established the genus Nevrorthus and -in describing N. iridipennis -provided the first (and quite wondrous) illustration of a nevrorthid (Fig. 1). In the opus magnum of Anton Handlirsch "Die fossilen Insekten und die Phylogenie der rezenten Formen" (1906)(1907)(1908), Nevrorthidae were still cryptic and hidden within Sisyridae (as Sisyra (Rophalis) relicta Hagen, 1856). Krüger (1923) treated the genera Rhophalis (sic) Erichson (sic) and Neurorthus (sic) Costa again as belonging to Sisyridae. They remained hidden in the phylogenetic tree of the Neuroptera by Withycombe (1925). In the meantime Nakahara (1915) erected the tribe Neurorthini, yet placed it in the Hemerobiidae: Hemerobiinae. Forty years later he raised Neurorthini to the subfamily level (Nakahara 1958), yet retained them within Sisyridae.
Zwick (1967) finally awarded family status to Neurorthinae Nakahara: Neurorthidae Nakahara, 1915, and discussed a sister group relationship of Neurorthidae with Osmylidae. Oswald and Penny (1991) re-established Nevrorthus Costa, 1863, as the clear intention of Costa and identified Neurorthus as a misspelling.
In two small and inconspicuous papers (U. Aspöck 1992Aspöck , 1993, Nevrorthidae received special phylogenetic attention and the following hypothesis was proposed: The Nevrorthidae do not belong to the Hemerobiformia as hitherto assumed but are interpreted as the sister group of the Myrmeleontiformia with a special head cervix articulation serving as a larval synapomorphy. In the first computerized analysis of the Neuropterida (U. Aspöck et al. 2001), the Nevrorthiformia emerged as sister group of Hemerobiformia + Myrmeleontiformia (with larval cryptonephry as a synapomorphy). The com-pact larval head capsule of Nevrorthiformia represents a basic pattern, the compact head capsule is retained in Myrmeleontiformia, where however, it is strongly modified, as emphasized in U. Aspöck (2002).
In the first molecular analysis of Neuropterida (Haring and U. Aspöck 2004), Nevrorthidae held their position as sister group of all other families, however, the Hemerobiformia were disrupted.
To escape the conflicting results, the phylogenetic relevance of the genital sclerites was tested on the basis of the gonocoxite concept put forward by U. Aspöck and H. Aspöck (2008a). In this analysis Nevrorthidae retained their position as sister group to all other families.
In the course of further molecular analyses, mentioned above, Nevrorthidae was retrieved either as a sister group to Sisyridae and Osmylidae and all three constituted a monophylum (Winterton et al. 2010), or Nevrorthidae and Sisyridae alone constituted the sister group to the rest of Neuroptera except Coniopterygidae (Wang et al. 2016).
In context of a microcomputer analysis of the larval head of Nevrorthus (Beutel et al. 2010), the sister group relationship of Megaloptera + Neuroptera was corroborated, and Nevrorthidae was confirmed as sister group of Myrmeleontiformia + (the reinstalled) Hemerobiformia. In the context of that analysis, three-dimensional reconstructions of the larval head not only of Nevrorthus but also previous ones concerning Raphidioptera (Beutel and Ge 2008) and Megaloptera (Beutel and Friedrich 2008) became essential for better understanding the evolution of the characters. A special focus on the head morphology of adult Neuroptera by Zimmermann et al. (2011) and Randolf et al. (2013Randolf et al. ( , 2014 ended up with Sisyridae as sistergroup of all other Neuroptera, followed by Nevrorthidae as sistergroup of the remaining families. The discovery of the mouthpart muscle M. stipitalis transversalis and a hypopharyngeal transverse ligament found in the head of N. apatelios was newly discovered for Neuroptera and herewith for the first time in Endopterygota by Randolf et al. (2014). In addition a submental gland with multiporous opening -apparently unique among insects -was described for Nevrorthidae and Osmylidae (Randolf et al. 2014). The phylogenetic relevance of the adult head in Nevrorthidae is obvious.

Biology
The unique aquatic larva of Nevrorthus fallax was discovered and described in detail by Zwick (1967). The first description of a nevrorthid larva, however, is much older (Takahashi 1942), but it was subordinated under Dilaridae. Larvae are carnivorous and live in the coarsely granular sands of clear, clean rivulets. Pupation takes place in the water on the undersides of stones. The silky cocoon spun by the larva comprises two layers (Malicky 1984). An aquatic pupa is unique among Neuroptera and Neuropterida. The length of development has not been adequately investigated. Probably, it takes one year. Nothing is known on the number of eggs laid by a female. Adults (Fig. 2) are found on leaves of overhanging tree branches and on bushes and low vegetation close to the water. They are active in the day-time and are rarely attracted by artificial light. Malicky (1984) found adults on sticky leaves and assumed honeydew to be an important part of the diet. The adaptations of the mouthparts, namely paraglossae that are folded onto the ligula thus forming a secondary prolongation of the salivary opening to the tip of the ligula (Randolf et al. 2013(Randolf et al. , 2014) are interpreted as adaptations to feeding not only on liquid but also on desiccated honeydew. A further indication for glycophagous feeding habit has been studied already by Kokubu and Duelli (1983). Monserrat (2005) found fungal spores in the digestive tract of Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel, 1977, and Nipponeurorthus fasciatus Nakahara, 1958. Randolf et al. (2014 described the mouthparts of Nevrorthus apatelios as instruments with strongly sclerotized asymmetrical mandibles with apical incisors which indicate a carnivorous feeding habit (Stelzl 1992).

Fossil taxa
At present, fossil Nevrorthidae have been found in Eocene Baltic amber (about 45 million years BP) and in mid-Cretaceous Burmese amber (about 100 million years BP, species with familial placement not confirmed and undescribed).
As concerns fossil Nevrorthidae, all available knowledge of material from the Baltic amber has been summarized recently (Wichard 2016). The excellent preservation especially of the genital sclerites of most species allows homologisation with extant species, which is fascinating. However, the Baltic amber material is too young to interpret deeper phylogeny. This may also be the case with the much older Burmese amber (Grimaldi et al. 2002, Xia et al. 2015, from which more surprising findings are to be expected. Further information on fossil Nevrorthidae can be found in Berendt (1845Berendt ( -1856, Nel and Jarzembowski (1997), Makarkin and Perkovsky (2009), Wichard et al. (2009, Wedmann et al. (2013), Wichard (2014Wichard ( , 2016, Makarkin (2016).
The photograph of the larva was made with a Canon EOS 350D (Fig. 1b). Stacked digital images (Figs 1, 3, 4a, f-h) were taken with a Leica DFC camera attached to a Leica MZ16 binocular microscope and processed with the help of Leica Application Suite. They were then stacked with Zerene Stacker 64-bit and processed with Adobe Photoshop Elements 8. Other images (Figs 4b-e, i-k) were made with a Nikon D800 attached with a Nikon AF Micro-NIKKOR 105mm f/2.8D lens.

Illustrations
Genitalic preparations in connection with redescriptions were made by clearing the apex of the abdomen in a cold saturated KOH solution for 3 h. After rinsing the KOH with acetic acid and water, the apex of the abdomen was transferred to glycerine for further dissection and examination. Drawings of the genitalia were made with a camera lucida of a Leica WILD M 10 at the NHMW and with a Leica S8 APO at the CAU. The genital structures were interpreted and labelled on the basis of the gonocoxite-concept hypothesized by U. Aspöck and H. Aspöck (2008a, b).
Distribution maps were provided with ArcMap ver. 10.3.1.4959 based on the distribution records provided in the Supplementary material 1. Source of the maps: National Geographic-Weltkarte -Content may not reflect National Geographic's current map policy. Sources: National Geographic, Esri, DeLorme, HERE, UNEP-WC-MC, USGS, NASA, ESA, METI, NRCAN, GEBCO, NOAA, increment P Corp.

Redescriptions
In the redescriptions of the species the homology hypotheses and the terminology of the genital sclerites developed by U. Aspöck and H. Aspöck (2008a, b) are applied.

Character description and phylogenetic analysis
The present phylogenetic analysis aimed to reconstruct the intergeneric phylogeny of Nevrorthidae. Morphological characters were used for the phylogenetic inference. Thirty-one characters were coded with 27 binary and four multistate (see Supplementary material 2). The character matrix can be found in Supplementary material 3. All characters were treated as unordered and with equal weight. The multistate characters were treated as additive. Italoraphidia solariana (Navás, 1928) (Raphidioptera: Raphidiidae) and Megalomus tortricoides Rambur, 1842 (Neuroptera: Hemerobiidae) were selected as the outgroup taxa. The ingroup taxa include all extant and fossil species of Nevrorthidae previously described. However, two species of Nipponeurorthus (i.e., Ni. qinicus and Ni. tinctipennis) and one species of Proberotha (i.e., P. dichotoma), which lack a large amount of data, were excluded for an additional analysis. Analyses were performed using NONA ver. 2.0 (Goloboff 1993) with a heuristic search. Maximum number of trees to keep was set to 10000 and the number of replications to 100. The bootstrap branch support values were calculated in NONA ver. 2.0. Character states were mapped on the strict consensus tree using WinClada ver. 1.00.08 (Nixon 2002), showing only unambiguous changes.
Nevrorthidae are alternately addressed as enigmatic or mysterious -but why? The adults look rather inconspicuous and may even be frequent if one searches for them at the right place and at the right time. Even the cryptic larvae, which inhabit sandy and stony grounds of rivulets may be frequent if one searches for them at the right place and at the right time. The aquatic pupae are certainly unique among Neuropterida, but neither enigmatic nor mysterious. The secret around the mystery concerning Nevrorthidae may be their isolated existence in hidden mountainous rivulets and the hypothesis that there are hitherto undiscovered remote relic places harbouring populations of known or still unknown species. Male. Body length 2.2 mm; forewing length 6.0-7.5 mm, hindwing length 5.5-6.5 mm.
Biology and ecology. Adults have been taken from May to October, most specimens were collected in June and July. The known vertical distribution is 90-1400 m. The larva is found in mountain rivers (the temperature of inhabited brooks varied from 11.9-21.5°C).
Head yellowish, dark brown line at middle. Antennae pale yellow, scapus and pedicellus brownish. Mouthparts yellow.
Biology and ecology. Adults have been taken from March-October, most specimens were collected in June. The known vertical distribution is 70-1050 m. The larva is known and has been described (Zwick 1967). Larvae inhabit the stony bottom of cold (5-10°C) swiftly running mountain brooks (Zwick 1967), We have, however, found the species in Sardinia also at the estuary of a river a few meters above sea level. Malicky (1984)  Abdomen dorsally brown with yellow pattern, ventrally yellowish with only a few brownish spots. Gonocoxites 9 as huge plates, gonostyli 9 digitiform, gonapophyses 9 processus-like; ectoproct broadly rounded. Complex of gonocoxites + gonostyli + gonapophyses 10 amalgamated with sternite 9, forming a pseudoapex of the latter and framing it laterally, terminally with short incision. Gonocoxites 11 fused into a bow-like bridge.
Biology and ecology. Adults have been taken from April-June; most specimens were collected in May. The known vertical distribution is 336-530 m. Larvae were found in small brooks. Temperature of inhabited brooks varied from 13.6-15.7°C (Malicky 1984).
Specimens examined and records published. Supplementary material 1. Lectotype female (by explicit designation): Calabria, Reggio Calabria "Valli di Aspromonte" (MZUN), Pantaleoni (designated 1993Pantaleoni (designated , published 1999. Biology and ecology. Adults have been taken from May-July; most specimens were collected in May. The known vertical distribution is 354-1350 m. The larva is known and has been described (Malicky 1984), the temperature of inhabited brooks measured varied from 7.9-23.8°C.
Biology and ecology. Adults have been taken from November-February. There is no data concerning the vertical distribution. The larva of A. brunneipennis is possibly known, however, it cannot be differentiated from that of A. horstaspoecki (see Austroneurorthus sp. in Fig. 15).
Biology and ecology. Adults have been taken from December-February, with most specimens collected in February. There is no data concerning the vertical distribution. The larva of A. horstaspoecki is possibly known, however, it cannot be differentiated from that of A. brunneipennis (see the distribution of Austroneurorthus sp. in Fig. 15).

Diagnosis.
Adults of small body-size; male forewing length 6-10 mm. Body coloration generally yellow. Fore-wings transparent to pale yellowish brown, sometimes with brown markings, sometimes with spectacular colour pattern. Costal crossveins of forewings at least partially forked in most species. Hindwing MA and anterior branch of MP forked distal to outer series of gradate crossveins in most species. Male abdominal segment 7 sometimes enlarged. A ring-like zone of glands sometimes present between male abdominal segments 8 and 9. Abdominal eversible sacks -as e.g. in Nevrorthus -are so far found only in Nipponeurorthus fasciatus (between segments 8 and 9). Male sternite 9 short, not strongly extending posteriad; gonocoxites 9 present as a pair of robust claspers, terminally with gonostyli 9; complex of gonocoxites + gonostyli + gonapophyses 10 present as a pair of discrete sclerites with long blade-like, spinous, or claw-like distal lobes, free or more or less attached (or amalgamated respectively) with sternite 9, as lateral "frame" and terminal sclerites (appearing as a pseudoapex of sternite 9); gonocoxites 11 reduced to sclerite claspers which might represent the gonostyli 11, located between bases of gonocoxites 9. Fused female gonocoxites 8 broad, nearly twice as long as tergite 8; gonocoxites 9 foliate or club-shaped; bursa copulatrix comprising a sclerotized structure. Distribution. China, Japan. Thorax yellow. Legs yellow; coxae, trochanter and femora slightly paler. Wings slightly yellowish brown, with pterostigmatic areas creamy yellow; forewing with distal margin brown and with distinct brown markings on gradate crossveins as well as on 1r-rs; other less distinct brown markings present on distal branching points of most longitudinal veins. Veins yellowish brown except for those in dark markings brown. Hindwing much paler than forewing, with distal dark edging much shorter and paler than that on forewing. Veins pale yellow, with 1r-rs and 2r-rs brown.

Nipponeurorthus damingshanicus
Abdomen yellow, dorsally largely tinged with pale reddish brown. Gonocoxite 9 robust on proximal half and strongly incurved on distal half, with a small hairy tubercle on inner surface; gonostylus 9 terminally flattened and bearing a spinous lobe. Ectoproct broad, directed posteroventrad, and concaved medially on posterior margin, with median portion slightly domed dorsad in lateral view, and with posterolateral corner protruding into a digitiform process. Complex of gonocoxites + gonostyli + gonapophyses 10 proximally broad, bearing a roundly tapered dorsal lobe and slender ventral lobe, distally with a long and blade-like projection; distal projections crossing each other at mid-length. Gonocoxite 11 not visible; gonostyli 11 present as posteriorly bifurcated sclerite.
Specimens examined and records published. Thorax yellow; pronotum with lateral margins slightly darker; meso-and metanota laterally with a pair of broad brown markings. Legs yellow, with 5th tarsomere slightly darker. Wings slightly yellowish brown, with pterostigmatic areas pale brown; forewing with distal and posterior margins almost brown and with pale brown markings on gradate crossveins as well as on 1r-rs; other pale brown markings present on branching points of most longitudinal veins. Veins yellowish brown except for those in dark markings brown. Hindwing much paler than forewing, with distal margin brown. Veins pale yellowish brown, with 1r-rs, 2r-rs, and gradate crossveins brown.
Fused gonocoxites 8 about 1.5 times as long as tergite 8, flatly plate-like. Gonapophyses 8 subtrapezoidal, largely covered by gonocoxite 8, lateral margins distinctly sclerotized. Bursa copulatrix comprising a generally subglobal sac-like structure, which is nearly as long as tergite 8; proximal portion moderately sclerotized, lateral portion protruding into a pair of ovoid membranous lobes, which are acutely pointed dorsad.
Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.
Thorax yellow. Legs yellow. Wings transparent, immaculate, with pterostigmatic areas dark yellow. Veins yellow, with costal crossveins slightly darker.
Abdomen yellow. Gonocoxite 9 robust on proximal half, with a small hairy tubercle on inner surface; distal half strongly incurved, with an obtuse ventral lobe; gonostylus 9 spinous and forked at tip. Ectoproct broad, directed posteriorly. Complex of gonocoxites + gonostyli + gonapophyses 10 rather small; lateral arms much longer than distal projections, strongly sinuate, and distinctly widened posteriorly; distal projections slenderly digitiform, rather close to each other, each projection laterally with a feebly sclerotized flat lobe. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 as posteriorly bifurcated sclerite.
Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.
Thorax yellow, with yellowish setae. Legs yellow throughout, with yellowish setae. Wings slightly yellowish brown, with pterostigmatic areas yellowish brown; forewing with distal margin brown, and with distinct brown markings on gradate crossveins as well as on 1rrs; other less distinct brown markings present on distal branching points of most longitudinal veins; veins yellowish brown except for those in dark markings brown; hindwing much paler than forewing, with distal dark edging much shorter and paler than that on forewing; veins pale yellow, with 1r-rs and 2r-rs brown.
Abdomen yellow. Gonocoxite 9 robust on proximal half and strongly incurved on distal half, ventrally with an upcurved short lobe separated from the main body of gonocoxite 9; inner surface with a small hairy tubercle; gonostylus 9 terminally rounded and bearing a spinous lobe. Ectoproct broad, directed posteriad, and subtrapezoidal and slightly concaved on posterior margin in dorsal view. Complex of gonocoxites + gonostyli + gonapophyses 10 proximally robust, distally with a slenderly spinous projection, which laterally bears a feebly sclerotized flat lobe. Gonocoxite 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as a posteriorly bifurcated sclerite. Hypandrium internum not visible. Biology and ecology. Adults have been taken in July. The known vertical distribution is 1600 m. The larva is unknown.
Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.
Thorax yellow. Legs yellow. Wings transparent, immaculate, with pterostigmatic areas yellow; longitudinal veins mostly yellow, except for those posterior to 2nd gradate crossveins brown; crossveins mostly brown, except for those on pterostigmatic areas yellow.
Abdomen yellow, dorsally much darker. Gonocoxite 9 robust on proximal half, with a small hairy tubercle on inner surface; distal half strongly incurved and sinuate, ventrally with two obtuse lobes, one directed outward and bald, the other directed inward and setose; gonostylus 9 acutely pointed but unforked. Ectoproct broad, directed posteroventrad, with posterior margin slightly concave. Complex of gonocoxites + gonostyli + gonapophyses 10 with lateral arms much longer than distal projections, straightly directed; distal projections digitiform, acutely pointed at tip, widely separated and parallelly directed with each other. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as posteriorly bifurcated sclerite.
Specimens examined and records published. Supplementary material 1. Syntypes: "Mt. Atago near Kyoto on July 2, '14" [A lectotype should be designated, however, the syntypes are currently unavailable and possibly even lost].
Biology and ecology. Adults have been taken from July-August. The known vertical distribution is 235-1000 m.
Distribution. Japan (Hokkaido, Honshu). Prothorax yellow, meso-and metathorax pale brown. Legs yellow. Wings transparent, with pterostigmatic areas pale yellow. Forewing with brown stripes along longitudinal veins posterior to 1st gradate crossveins and branches of CuA, CuP and 1A, and also with brown stripes on most crossveins except for those on pterostigmatic areas. Hindwing only with brownish stripes on 1r-rs and 2r-rs. Veins blackish brown on forewings and pale brown on hindwings, but costal crossveins on pterostigmatic areas and longitudinal veins on proximal half yellow.
Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.
Thorax yellow. Legs yellow. Wings transparent, immaculate, with pterostigmatic areas yellow; longitudinal veins yellow; crossveins mostly dark brown, except for those on pterostigmatic areas yellow.
Abdomen yellow, dorsally purplish brown. Gonocoxite 9 robust on proximal half, with a small hairy tubercle on inner surface; distal half strongly incurved; gonostylus 9 spinous and unforked. Ectoproct broad, directed posteroventrad, with posterior margin slightly concaved, and with a pair of subtriangular ventral projection. Complex of gonocoxites + gonostyli + gonapophyses10 transversely broad; lateral arms nearly as long as distal projections, arcuate, medially with a pair of projections, which are straightly directed dorsad and widened on distal half; distal projections digitiform, straightly and parallelly directed dorsad with each other. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as posteriorly bifurcated sclerite.
Biology and ecology. Adults have been taken from May-July, most specimens were collected in July. No data concerning the vertical distribution are available. The larva is unknown, however, Nakahara (1958) hypothesized an aquatic life style of the larva due to the findings of adults along rivers and brooks.
Distribution. Japan (Hokkaido, Honshu, Kyushu, Tsushima Island). Thorax yellow. Legs yellow. Wings transparent, with pterostigmatic areas pale yellow; forewing with brownish stripes on most crossveins except for costal crossveins and with brownish spots on distal branching points of most longitudinal vein; hindwing with brownish spots on distal branching points of Rs, MA and MP; veins mostly yellow, except for those on dark markings brown; costal crossveins on proximal half of forewing costal areas pale brown.
Fused gonocoxites 8 about 2.0 times as long as tergite 8, flatly plate-like. Gonapophyses 8 subtriangular, largely covered by gonocoxite 8, lateral margins distinctly sclerotized. Bursa copulatrix sac-like, suboval, much longer than tergite 8; proximal portion with a pair of broad sclerotized areas, median portion ventrally with a pair of sclerotized holes, distal portion marginally sclerotized and terminally curved dorsad in lateral view.
Specimens examined and records published. Supplementary material 1. Lectotype designation presently not possible (see above).
Biology and ecology. Adults have been taken from July-August, most specimens were collected in July. No data concerning the vertical distribution are available. The larva is unknown.
Thorax yellow. Legs yellow. Wings transparent, immaculate; veins mostly pale brown on forewings, except for veins on wing base and proximal half of anterior branch of MP yellow; veins mostly pale brown on hindwings, except for veins on wing base yellow.
Female. Unknown. Specimens examined and records published. Supplementary material 1. Holotype (by implicit monotypy) male: China, "Shaanxi, Ankang" (CAU). So far, the holotype has not been found in the entomological collection of CAU. There is a possibility that the primary type is lost or damaged. However, due to the lack of additional specimens of this species, we cannot designate a neotype.
Biology and ecology. No data are available. The larva is unknown.

Nipponeurorthus tianmushanus Yang & Gao, 2001
Wings slightly yellowish brown, with pterostigmatic areas pale brown; forewing with distal margin brown and with brownish markings on most crossveins except for costal crossveins; hindwing similarly patterned; veins pale brown.
Gonocoxite 9 robust on proximal half and strongly incurved on distal half. Ectoproct broad, directed posteroventrad, and strongly concaved on posterior margin. Complex of gonocoxites + gonostyli + gonapophyses 10 present as a pair of slender lobes, which are rather close to each other at the tip.
Female. Unknown. Specimens examined and records published. Supplementary material 1. Holotype (by original designation), male, China, "Zhejiang, Tianmushan, 22.VII.1963, Io Chou" (CAU). Thus far, the holotype has not been found in the entomological collection of CAU. There is a possibility that the primary type is lost or damaged. However, due to a lack of any additional specimens of this species, we cannot designate a neotype.
Biology and ecology. No data available. The larva is unknown.
Distribution. China (Zhejiang). Thorax yellow; meso-and metanota laterally much darker. Legs yellow. Wings transparent, immaculate, with pterostigmatic areas pale yellow; veins mostly yellowish brown, with crossveins much darker, and with C, Sc and R pale yellow on forewings; veins mostly pale yellow, with longitudinal veins of distal half and some crossveins (i.e. 1r-rs, 2r-rs, and gradate crossveins) pale brown on hindwings.
Biology and ecology. The adult has been taken in July. The known vertical distribution is 1800 m. The larva is unknown.
Head reddish orange, slightly shiny. Antennae blackish brown, with scape and pedicel pale yellowish brown, and with proximal two segments of flagellum orange. Mouthparts orange.
Thorax reddish orange, slightly shiny. Legs orange. Wings smoky brown, with slightly leathery membrane; veins blackish brown, with proximal half of C and extreme bases of other longitudinal veins much paler.
Pterostigmatic areas very dark, with their crossveins rather weak and obscure; Rs proximally 2-branched, both branches deeply bifurcated, with bifurcation nearly 1/2 as long as whole wing; all main branches having additional branching, terminally leaving 8-10 small bifurcate or trifurcate forks; MA completely fused with Rs proximally in forewing, but visible as an independent vein at base of hindwing; medially bifurcated, with both branches having additional branching, terminally leaving 8 small bifurcated or trifurcated forks; MP proximally 2-branched, each branch bifurcated at distal 1/3 in forewing and at distal 1/4 in hindwing, terminally leaving 8-10 small bifurcate or trifurcate forks; CuA 7 to 8-branched in forewings, terminally leaving ca. 10 small bifurcate or trifurcate forks, and 11 to 13-branched in hindwings, with proximal branches vertical to stem of CuA, terminally leaving 14-15 small bifurcate or trifurcate forks; CuP with a small bifurcate fork terminally; 1A terminally 4 to 5-branched in forewings and 3-branched in hindwings; 2A 7-branched in forewings and 6 or 8-branched in hindwings; 3A simple.
Male Biology and ecology. The only adult has been taken in May in the vicinity of a waterfall. The known vertical distribution is 1715 m. The larva is unknown.

Phylogenetic analysis
The parsimony analysis of the primary matrix including all species of Nevrorthidae yielded 7712 most parsimonious trees (MPT) (length = 49, consistency index = 73, retention index = 93) and the strict consensus tree is shown in Supplementary material 4. The phylogeny was poorly resolved probably due to the inclusion of several ingroup taxa with a large number of missing data. The monophyly of only three genera with more than one species was recovered, including Austroneurorthus, Nevrorthus and Palaeoneurorthus. The latter two genera formed a sister group, and together with Rophalis they formed a monophylum.
The parsimony analysis of the refined dataset with deletion of two species of Nipponeurorthus (i.e., Ni. qinicus and Ni. tinctipennis) and one species of Proberotha (i.e., P. dichotoma) yielded 40 most parsimonious trees (MPT) (length = 49, consistency index = 73, retention index = 92) and the strict consensus tree is shown in Figure  18. Based on these results, all Nipponeurorthus species formed a monophylum, supported by the male gonocoxite 9 with subdistal inflation and additional lobes (char. 18:1) and the female fused gonocoxites 8 much longer than wide with posterior tapering (char. 29:2). The monophyletic group comprising Rophalis, Nevrorthus and Palaeoneurorthus, which was recovered in the analysis of the primary dataset, was also recovered here and supported by the male gonocoxite 9 ventrally with a long lobe (char. 19:1) and the elongated male gonapophyses 9 with acute projections (char. 23:3). This monophyletic clade of three genera was grouped with Austroneurorthus and Electroneurorthus. The synapomorphic characters of the monophyletic group comprising Austroneurorthus, Electroneurorthus, Rophalis, Nevrorthus and Palaeoneurorthus include the elongated and posterodorsally directed male sternite 9 (char. 14:1 and char. 17:1), the ovoid male gonapophyses with several spines (char. 23:2), and the presence of fused gonocoxites 10 (char. 25:1). The phylogenetic positions of Balticoneurorthus, Proberotha and Sinoneurorthus were not resolved.

Phylogenetic position of Nevrorthidae
Irrespective of the fact that Nevrorthidae was assigned at various positions in different analyses based on morphological and molecular data (U. Aspöck et al. 2001, Haring and U. Aspöck 2004, U. Aspöck and H. Aspöck 2008a, Beutel et al. 2010, Winterton et al. 2010, Zimmermann et al. 2011, Randolf et al. 2013, Randolf et al. 2014, Wang et al. 2016, several hypotheses, which have been catalysed via Nevrorthidae, are of general significance regarding Neuropterida: The hypothesis of aquatic larvae as a synapomorphy of Megaloptera + Neuroptera induces the hypothesis that cryptonephry might be an answer to secondary terrestrial life-style of the crown clade within Neuroptera. Gaumont (1976) provided comparative studies of the sucking tubes, guts and the Malpighian tubules of Neuropteran larvae. In this connection she studied the phenomenon of cryptonephry of terrestrial larvae. She interpreted the free Malpighian tubules of aquatic larvae of Sisyridae and Nevrorthidae as secondary adaptations. We interpret free Malpighian tubules -at least in Nevrorthidae -as the plesiomorphic condition and the phenomenon of cryptonephry (= complex connection of the Malpighian tubules with the colon) as an adaptation to secondary terrestrial life style of the remaining families (U. Aspöck et al. 2001).
A compact head capsule with a large gula is interpreted as belonging to a ground pattern in larval Neuropterida. In Neuroptera this feature is retained only in Nevrorthidae, thus placing them in a key position within the order. An open or compact head capsule in connection with a loss of the gula (U. Aspöck and H. Aspöck 2010b) represent phylogenetic trends in the remaining Neuroptera (U. Aspöck and H. Aspöck 2007).
A neck-like, somewhat articulating cervix is apomorphic and a larval synapomorphy of Neuroptera. Several families (former Hemerobiformia) have lost this condition (U. Aspöck et al. 2001). The region underwent further elongation in Nevrorthidae and is known as the socalled "Rollengelenk" (Zwick 1967).
Pleuritocavae, paired sacks of uncertain, possibly pheromonal, function -a curiosity of male adults -have been found ventrally between segments 6 and 7 in Nevrorthus (U. Aspöck and H. Aspöck 1983) and R. relicta (Wichard et al. 2009), between segments 7 and 8 in R. relicta, between segments 8 and 9 in Ni. fuscinervis, Ni. multilineatus and R. relicta, and dorsally between tergites 8 and 9 in Ni. fasciatus and R. relicta. These sacks are only visible when they are everted, so they are possibly more common than previously suspected. Similar structures are found in other Neuroptera, especially Nemopteridae. A phylogenetic relevance may be assigned to them, however, the character is unreliable due to the variable pheromonal status of the observed individual specimens.
A most recent study on mitochondrial phylogenomics of the Neuropterida (Wang et al. 2016) corroborates a sister group relationship of Megaloptera + Neuroptera and a sister group relationship of Coniopterygidae + monophyletic remaining Neuroptera. Within the Neuropteran families excluding Coniopterygidae, the clade Sisyridae + Nevrorthidae was assigned as sister group to Osmylidae + the monophylum constituted by the remaining twelve families. The sister group relationship of Nevrorthidae + Sisyridae has been discussed in detail in Wang et al. (2016) especially with respect to the morphological disparity of the larvae of the two families. This ongoing discussion remains a challenge in our understanding of Nevrorthidae.

Intergeneric phylogeny within Nevrorthidae
By sharing a number of apomorphic characters, among the four extant genera of Nevrorthidae, it is not difficult to infer a close relationship between Austroneurorthus and Nevrorthus. The phylogenetic position of Sinoneurorthus is still unclear due to the lack of male specimens, yet it appears to be similar to Nipponeurorthus by having the partially branched forewing costal crossveins and similar modification of bursa copulatrix. Based on the presently reconstructed phylogeny, the Eocene Baltic amber genera of Nevrorthidae appear to be heterogeneous. Electroneurorthus, Rophalis and Palaeoneurorthus were assigned in the same clade with the extant Austroneurorthus and Nevrorthus. Balticoneurorthus and Proberotha have unresolved phylogenetic positions, while they seem to be relatively basal groups having few apomorphic characters. Alternatively, they might be closely related to Nipponeurorthus by having the partially forked forewing costal crossveins and the similar male gonocoxites 9.
The most interesting discovery in connection with nevrorthid genital sclerites is the complex constituted by the gonocoxites, gonostyli and gonapophyses of segment 10, which is discernible, e.g. in Ni. pallidinervis on one hand, but completely camouflaged in all Nevrorthus species on the other hand. In these species it appears as an elongated apex (pseudoapex) of sternite 9. This phenomenon in Nevrorthidae plays a key role in the homologisation of the genital sclerites based on the gonocoxite concept developed in U. Aspöck and H. Aspöck (2008a) which draws upon the hypothesis of traceable gonocoxites, gonostyli and gonapophyses in segment 9, as well as in segments 10 and 11, irrespective of the fact that these segments are highly transformed in connection with their functions in copulation. Additionally, the modifications of these sclerites are important for inferring the intergeneric phylogeny of Nevrorthidae. Moreover, a ring of glands between segments 7 and 8 in males of Nevrorthus, between segments 8 and 9 in males of Austroneurorthus and several species of Nipponeurorthus seems to be a more authentic character since it is apparently more stable than the eversible sacks. The feature may have phylogenetic relevance; however, it cannot be traced reliably in fossil specimens.

Biogeography
The world distribution of Nevrorthidae demonstrates the relictual nature of this family. They are "living fossils" in the sense of Thenius (2000) for several reasons -the disjunct distribution, low number of extant species and the archaic shape of the larval head capsule. Although the number of fossils of Nevrorthidae is continuously growing, those known from the Eocene Baltic amber, as well as from the mid-Cretaceous Burmese amber, provide limited evidence to understand the present-day disjunctive pattern. Their characterisation as faunal elements with respect to glacial refugial centres in the sense of de Lattin (1967)  Questions to be asked concern quite different phenomena.
Why are there no Nevrorthidae either in Nearctic and Neotropical regions or the Afrotropics?
The recently discovered N. reconditus answers our old perpetuating question as to why Nevrorthidae are absent in the western Mediterranean -because they are already there! Nonetheless, the question why the genus Nevrorthus is lacking in the eastern Mediterranean, still remains. Present climate change: Recent findings of N. apatelios in the Alpine regions of Friuli and Slovenia represent the northernmost records of the family in Europe, thus making it a Central European matter, triggering further hypotheses on the distribution of this puzzling family. Have Nevrorthidae been continuously overlooked north of the Alps? Ceratinly not! Aquatic insects are in general well explored -new discoveries as the above mentioned are therefore more than surprising. Most probably N. apatelios reached Friuli from rivers in northern Italy and survived the last glacial period in extramediterranean-European refugial centres south of the Alps (U. Aspöck and H. Aspöck 2010a).
The surprising discovery of the spectacular Sinoneurorthus yunnanicus (Liu et al. 2012) in China and the continuous discovery of new nevrorthid species in eastern Asia (Liu et al. 2014) denote this part of the world as a hot-spot of nevrorthid evolution. These recent findings of Nevrorthidae in mainland China weaken our previous hypothesis that Austroneurorthus, and partly also Nipponeurorthus, show a coastal distribution pattern (the socalled Tethys distribution pattern) (Starmühlner 1982, U. Aspöck 2004. It becomes clear that some taxa occur far from the sea. Based on male genitalia, Nevrorthus is the sister group of Austroneurorthus -however, biogeographically this infers a severe conflict. lege Station, Texas), Akihiko Shinohara and Utsugi Jinbo (Tokyo), Fumio Hayashi (Tokyo), Günther Theischinger (Sydney), and Peter Zwick (Schlitz) for providing material and for their patience with overdue loans. Harald Bruckner (Vienna) is gratefully acknowledged for taking many of the photographs, arranging the figures, providing the list of material for the Supplementary materials and preparing the distribution maps. Many thanks to Franziska Denner (former Anderle) and to Peter Sehnal (Vienna) for taking photographs of living specimens. A big thank you goes to Silke Schweiger (Vienna) for helping with the logistics of these maps. Cordial thanks also to Eva Hitzinger for various technical assistances. Sincere thanks to Dušan Devetak (Maribor), Alexi Popv (Sofia) and to Susanne Randolf (Vienna) for thoroughly reviewing and improving the manuscript. Grateful thanks to Dr. John Plant (Guilford, Connecticut) for critically reading the manuscript and for polishing the English. The present study was funded by the National Natural Science Foundation of China (Nos. 31672322, 31322051, 41271063) (Rohdendorf 1938, Mostovski 1996a,b, 1997, J Zhang and H Zhang 2003, K Zhang et al. 2007a,b, 2008, 2009, 2010a,b, J Zhang 2010, 2012a,b, 2015, Oberprieler and Yeates 2012, Wang et al. 2017). An updated list of all the archisargid species is presented herein (see Table 1). Among them, the placement of Helempis yixianensis Ren, 1998 has been transformed. A recently erected species Archirhagio gracilentus Wang et al., 2017 and a new species Flagellisargus (Changbingisargus) parvus sp. n. described below are also supplemented (see Table 1).

Material and methods
Material. The specimens of shale fossil impression of a male and a female archisargid flies described herein are deposited in the collections of the Nanjing Institute of Geology and Palaeontology (NIGP), the Chinese Academy Illustrations. Specimen descriptions, photomicrographs and line drawings were done without immersion with the exception of photographs of details of the antennae and tibial spurs. The line drawings were produced with the aid of a camera lucida and the digital photomicrographs were taken using a stereomicroscope.
Colour described here refers to that of the fossil, where patterning is preserved.
Wing venation terminology follows that of Wootton and Ennos (1989) and Shcherbakov et al. (1995). The cell traditionally named the anal cell is, in fact, considered to be the cubital cell herein. Diagnosis. Small-size archisargid flies (body excluding antenna and genitalia less than 5 mm long); antennal scape long; arista (or stylus) absent; fork of R4+5 shallow, distad of level of R2+3 end; R5 ending before wing tip; discal cell short and wide (nearly 2.3 times as long as wide).
Distribution. Jurassic, China. Remarks. The subgenus Flagellisargus (Flagellisargus) stat. n. includes three known species: Flagellisargus (Flagellisargus) robustus J Zhang, 2012a, Flagellisargus (Flagellisargus) sinicus J Zhang, 2012a and Flagellisargus (Flagellisargus) venustus J Zhang, 2012a. Among them, the first and the third species are erected based on nearly complete male flies, the second one with head and abdomen missing. This known subgenus differs from Flagellisargus (Changbingisargus) subgen. n. in the following aspects: moderate-size archisargid flies (body excluding antenna and genitalia more than 9 mm long); antennal scape short; arista (or stylus) present; fork of R4+5 relatively deep, just at level of R2+3 end; R5 ending just at wing tip; discal cell narrow and long (nearly three times or more as long as wide).
Although the head and abdomen are missing, Flagellisargus (Flagellisargus) venustus demonstrates close similarities in wing venation to that of Flagellisargus (Flagellisargus) sinicus and Flagellisargus (Flagellisargus) robustus: fork of R4+5 relatively deep, just at level of R2+3 end; R5 ending at wing tip; discal cell narrow and long, nearly 3.5 times as long as wide (Fig. 3A-C). Thus, Flagellisargus (Flagellisargus) venustus should be retained in Flagellisargus (Flagellisargus) rather than be assigned to Flagellisargus (Changbingisargus) subgen. n. Diagnosis. Male archisargid flies 4.9 mm long (excluding antenna); antenna longer than head, scape more than one half of flagellum length; stem of Rs nearly as long as bR4+5; first fork of Rs slightly basad of level of M fork; crossvein r-m linking anterior margin of discal cell near to M fork; crossvein m-m long; section of mM3+4 short; male genitalia large, gonostylus subquadrate with apical denticle medially.
Abdomen with seven segments visible, nearly ovate-oblong, fourth widest, and nearly as wide as thorax, 1.8 times longer than head (excluding antenna) and thorax combined; genitalia rather large, subovate, longer but narrower than seventh abdominal segment, gonocoxite more or less oblong with its inner and outer margins slightly curved outwards, gonostylus subquadrate, wider than long, with a triangular apical denticle curved upwards, aedeagus invisible (Figs 1C, 2C).
Dimensions. Holotype (NIGP DHG 201701): length of body, 4.9 mm; head (excluding antenna), 0.7 mm; Diagnosis. Moderate-size archisargid flies (body excluding antenna and genitalia more than 9 mm long); antennal scape short; arista (or stylus) present; fork of R4+5 relatively deep, just at level of R2+3 end; R5 ending just at wing tip; discal cell narrow and long (nearly three times or more as long as wide).

Flagellisargus (Flagellisargus) cf. sinicus J. Zhang, 2012a
Abdomen with nine segments visible, nearly cylindrical, just a little narrower than thorax, 1.9 times longer than head (excluding antenna) and thorax combined; each of tergites with a wide, longitudinal, intermediate marking which is darkish brown; apex of abdomen with a scelerotized, needle-like ovipositor, and slightly longer than ninth segment (Fig. 4A).
Remarks. On the following characters, this fly could be assigned to Flagellisargus (Flagellisargus): body (excluding antenna and ovipositor) moderate-size (more than 9 mm long); antennal scape short (not elongated); arista (or stylus) well developed (about a quarter of flagellum length); fork of R4+5 just at level of R2+3 end; and R5 ending at wing tip.
Owing to having special characteristics (antennal flagellum with a darkish brown longitudinal furrow near to its outer margin and connecting base of arista and a tibial spur of hindleg well developed) this specimen shows close similarities in antennal and leg's structures to that of the known species Flagellisargus (Flagellisargus) sinicus. Unfortunately, its wing is incompletely preserved, and the discal cell, posterior branch of M, CuA, CuP and crossvein m-cu are rather ambiguous or invisible. For this reason, this impression fly could only be identified as Flagellisargus (Flagellisargus) cf. sinicus.

Discussion
Recently, Grimaldi and Barden (2016) proposed a single most-parsimonious tree indicating the relationships within the superfamily Archisargoidea. They considered that three genera possessing the plesiomorphic condition of unmodified (non aculeate) female terminalia are not basal to Archisargoidea: Daohugosargus J Zhang, 2012b, Orientosargus J Zhang, 2012band Uranorhagio K Zhang, 2010. Meanwhile, "Flagellisargus has a plesiomorphic, non stylate type of antenna and may also lie outside the Archisargoidea sensu stricto, but this would need to be confirmed with female specimens (only males presently are known)" (Grimaldi and Barden 2016: 17).
However, this study argues that Flagellisargus has a well developed arista (or stylus) although it is short. This crucial character had been illustrated in the original generic diagnosis and specific descriptions (J Zhang 2012a: 879, 881, Figs 3, 7). Furthermore, the female Flagellisargus has been discovered, and described herein. Flagellisargus (Flagellisargus) cf. sinicus has a scelerotized, needle-like ovipositor (Fig 4A). It is clear that Flagellisargus should be an arichsargid genus even according to the alternative classification proposed by Grimaldi and Barden (2016).
As for Daohugosargus, this genus was proposed for Sharasargus eximius K Zhang et al., 2008, which is a monotypic genus based on an incomplete impression fly with terminal abdominal segments missing (K Zhang et al. 2008). Its sex is uncertain. It is difficult to see how this genus could be distinguished as a female fly, let alone with unmodified (non aculeate) female terminalia. Daohugosargus demonstrates plesiomorphic similarities in body structures (as preserved) and in wing venation to those uncontested archisargids, and differs only from them by the characteristic vein R2+3 which is short, S-shaped, and arising late from Rs. It would be unreasonable to move this genus out of the superfamily Archisargoidea based only on this difference. This study considers that Daohugosargus is related rather to Archisargidae, Archisargoidea than to any other superfamilial groups.
Furthermore, the conclusion is debatable whether genera having non-aculeate female terminalia lie outside of Archisargoidea. For example, there are two species, Archirhagio striatus J Zhang et H Zhang, 2003 andArchirhagio varius J Zhang, 2015, belonging to the archisargid genus Archihagio Rohdendorf, 1938 that need consideration. The former species has a highly sclerotized, aculeate ovipositor; while, the latter one possesses a blunt, enlarged, fleshy, hook-like ovipositor (Wang et al. 2017: Figs 4D, E, originally the "ovipositor" was labelled as a "hypogynial valve"). However, Archirhagio varius cannot be excluded from Archihagio based on its species diagnosis although it has a non-aculeate female terminalia. Another example is the two species of Ovisargus Mostovski, 1996: O. gracilis Mostovski, 1996and O. singulus J Zhang, 2015. The former species has an aculeate ovipositor but the latter one has a podgy, conical (non aculeate) ovipositor. O. singulus should be assigned to Ovisargus based on the similarities in body structures and wing venation to that of O. gricilis regardless of the ovipositor. In addition, an aculeate ovipositor has evolved homoplastically in Diptera. It occurs in various groups, including a few Tipulidae, Phoridae, Pipunculidae, some Conopidae, Tephritoidea, Cryptochaetidae and Tachinidae (Pritchard 1983, Feener and Brown 1997, Skevington & Dang 2002, Stireman 2006, Grimaldi et al. 2011, Q Zhang et al. 2016. None of these groups (superfamilies or families) are distinguished based only on the specialized ovipositor. It is evident that the aculeate ovipositor is a convergent development in functional morphology, and does not reveal relationships between these taxa.
Using a geometric morphometric analysis, Wang et al. (2017) reviewed and revised the classification of Archirhagio. They redefined the diagnosis of Archirhagio zhangi K Zhang et al., 2009, andproposed Archirhagio mostovskii J Zhang, 2015 to be a junior synonym for Archirhagio zhangi based mainly on some similarities of wing venation and shape of abdominal segments. This study argues that both species were erected based on almost complete impression fossils of the male flies. As the placement is debatable, an overall, further comparative analysis in body structures and wing venation was necessary. Wang et al. (2017) ignored the sharp difference between both holotypes in some key taxonomic characteristics. Archirhagio mostovskii differs from Archirhagio zhangi in the following aspects: (1) male holoptic vs male dichoptic; (2) markings on abdominal tergites differ sharply; (3) size and shape of wing and wing venation differ distinctly; and (4) male genitalia differ distinctly. Thus, Archirhagio mostovskii can be separated from Archirhagio zhangi. Some detailed explanations are given as follows. In Diptera, the eyes of most families are holoptic (Cumming and Wood 2009); only a few families have a dichoptic male that is used in the family diagnosis in Lower Brachycera, e.g. Asilidae and Xylophigidae (Fisher 2009, Woodley 2009a. It is clear that the condition (male holoptic or dichoptic) is an important diagnosis for the identification of the Lower Brachycera. Both species, Archirhagio mostovskii and Archirhagio zhangi, are erected based on males, the former species having holoptic eyes with a very long midline (J Zhang 2015: Figs 2B, 4A); in contrast, the latter one has dichoptic eyes, which are widely separate (K Zhang et al. 2009 : Fig. 2). It is impossible that the different compound eye types of the male mentioned above occur in the same species. On the basis of these crucial taxonomic characters, Archirhagio mostovskii should be separated from Archirhagio zhangi.
Secondly, the shape and arrangement of the abdominal markings frequently provide useful taxonomic characters for dividing various groups of the Lower Brachycera, at least at species level, and many such studies have been published (Jones and Anthony 1964, Smith 1989, Woodley 2009a. Archirhagio zhangi shows each of abdominal tergites I-VI with a patch at the posterolateral corner (a left patch in segments II-IV is also present but badly preserved to judge from the original photomicrograph -see K Zhang et al. 2009: Fig. 1A, Fig. 5B herein). In contrast, Archirhagio mostovskii has a wide, medially longitudinal stripe and a wide transverse band along the  K Zhang et al., 2009(after K Zhang et al., 2009, C male terminalia (after K Zhang et al., 2009, modified). Scale bars 1mm.
hind margin on each of the abdominal tergites IV-VII, and the markings occupy almost the whole of tergites I-III (Fig. 5A herein). The sharply different markings on the abdominal tergites indicate that both male holotypes cannot be assigned to one and the same species.
Thirdly, Archirhagio mostovskii shows the wings are clearly shorter and wider than that of Archirhagio zhangi, (wing 12.1-13 mm long, 3.0-3.4 mm wide, about three times as long as wide vs wing 17.5 mm long, 3.8 mm wide, 4.6 times as long as wide); the wing is about one half of body length in the former species vs about fourth-fifths in the latter one. It should be pointed out that the revised diagnosis of Archirhagio zhangi defining body length between 29 and 32 mm, wing length between 20 and 23 mm, is questionable because the holotype of Archirhagio zhangi (body 21 mm long, wing 17.5 mm long) and the holotype of Archirhagio mostovskii (body 22.2 mm long, wing 12.1-13 mm long) falls distinctly short of that size. This revised diagnosis is related neither to Archirhagio zhangi nor to Archirhagio mostovskii. Furthermore, in wing venation the character of cell r1 closed or nearly so is an important diagnosis for Archirhagio mostovskii, differing from Archirhagio zhangi, in which cell r1 is clearly open. This crucial character demonstrates close similarity to that of Calosargus Mostovski, 1997, another archisargid genus. Nevertheless, in Calosargus the cell r1 is closed before the anterior margin of the wing, which has a very short or relatively long petiole apically [e.g. Calosargus (Pterosargus) sinicus J Zhang, 2010 and Calosargus (Pterosargus) thanasymus Mostovski, 1997]. This key character mainly differentiates Calosargus from Archirhagio. It is interesting that Archirhagio mostovskii is considered as a connecting link between Archirhagio and Calosargus. On balance, one should keep Archirhagio mostovskii as a separate species referred to Archirhagio but closely related to Calosargus.
Finally, the structural characteristics of male terminalia provide an unparalleled array of taxonomic characters in Diptera (McAlpine et al. 1981). "Male terminalia are a key morphological source of characters used to distinguish species in the vast majority of Diptera families and there are few modern taxonomic studies that do not include illustrations of male terminalia to aid in species diagnoses" (Sinclair et al. 2013). However, Wang et al. (2017) did not describe and illustrate the characteristics of male terminalia in the revised species diagnosis of Archirhagio zhangi, although they also commented that the original description of the male terminalia was incorrect. They only supplied two photomicrographs of an unnumbered specimen instead of the holotype male terminalia of Archirhagio zhangi (Wang et al. 2017: Figs 4B, C). Furthermore, they claimed that there are no significant modifications in male terminalia across the genus Archirhagio, consisting of the reduced ninth tergite, unsegmented gonocoxites, and pair of large parameres (Wang et al. 2017). Meanwhile, without providing any reference and citation, they declared that the terminology "aedeagus" used by J Zhang (2015) is incorrect, and should be instead of paired "parameres" (Wang et al. 2017). These deductions proposed by them clearly run counter to what many dipterists have concluded (McAlpine et al. 1981, Woodley 1989, Cumming and Wood 2009, Sinclair et al. 2013. This study argues that the kidney-shaped gonocoxite, bipectinate gonostylus and short and stout aedeagus demonstrate Archirhagio mostovskii as having distinctly different structures in the male terminalia from the specimens provided by Wang et al. (2017: Figs 4A,B,C). Unfortunately, there is neither description nor line drawing of the male terminalia of Archirhagio provided in their article; and thus a further comparison of male terminalia between Archirhagio mostovskii and Archirhagio zhangi is difficult herein. On the other hand, if those male terminalia investigated by them possess the same structures, then those specimens most likely belong to one and the same species that differs from Archirhagio mostovskii. It should be also noted that in Stratiomyomorpha + Muscomorpha sensu Woodley (1989), the aedeagus is indistinguishably fused to the parameral sheath to form the phallus (Cumming and Wood 2009). Currently, Archisargidae is assigned either within or near to the Stratiomyomorpha (Oberprielar and Yeates 2012) or Archisargoidea is probably an extinct sister group to the Muscomorpha (Grimaldi and Barden 2016). In any case, the paired parameres should be indistinguishable in the male terminalia of archisargids [e.g. Flagellisargus (Flagellisargus) sinicus -see J Zhang 2012a: Fig. 3] The so-called parameres of Arichrhagio, an undouble archisargid genus, identified by Wang et al. (2017) should be phallus "(aedeagus sensu authors concerning Stratiomyomorpha and Muscomorpha sensu Woodley, 1989)" (Cumming and Wood 2009).
Originally, the genus Helempis Ren, 1998 including two species: H. yixianensis and H. eucalla Ren, 1998 was placed in Protempididae (Ren 1998). The present author (J Zhang 2012b) thought that the two species could be united into one species, namely H. yixianensis, and Helempis, as a subgenus, could be transferred into Ovisargus referred to Archisardinae, Archisargidae. Through further contrastive studying, it could be reasonable to retain Helempis as a separate genus within the Archisarginae, Archisargidae. It differs from Ovisargus by the elongated discal cell and the deeper fork of R4+5, which is distinctly basad to R2+3 end.

Introduction
By the 1960s, it was revealed that the Balkan Peninsula was a distribution (and evolution) centre of the order Raphidioptera with an incredibly high number of spe-cies (H. Aspöck and U. Aspöck 1965). In the course of the following years, extensive field studies were carried out in various parts of the Balkan Peninsula (H. Aspöck 1987, H. Aspöck et al. 1989, H. Rausch and R. Rausch 2004. These investigations led to the discovery of nu- Dtsch. Entomol. Z. 64 (2) 2017, 123-131 | DOI 10.3897/dez.64.19859 Copyright Horst Aspöck et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. merous new species and among them a snakefly species, which -despite slight differences -was morphologically so similar to Raphidia ophiopsis Linnaeus, 1758, that we hesitated to separate it from R. ophiopsis. However, after discovery of unusually large populations of this taxon in various parts of Greece and in biotopes (e.g. in coastal areas with maquis vegetation) ecologically entirely different from those of R. ophiopsis, which is associated with coniferous trees, we decided to describe it as a subspecies of Raphidia ophiopsis: R. ophiopsis mediterranea (H. Aspöck et al. 1977). In the meantime, the taxon was surprisingly found in Italy (Apulia) and later in northwest Anatolia. In our monograph (H. Aspöck et al. 1991) we argued that the disjunct distribution could hardly be explained by natural dispersal and we therefore considered that human activities might have been a significant cause of the amplification of the distribution of R. o. mediterranea. One of our arguments was that R. o. mediterranea occurs on the eastern coast of the Apennine Peninsula around Brindisi, a region known for its intensive ship traffic with Greece, which dates back to antiquity. Subsequently, entomologists from Italy found R. o. mediterranea in western parts of the Apennine Peninsula, and from this they concluded that R. o. mediterranea had not been introduced from Greece to Italy by human activities, but that its occurrence in Italy was due to natural dispersal (Letardi 2002, Letardi and Pantaleoni 1996, Pantaleoni 2005. Meanwhile, the taxon was unexpectedly found in Romania (Kis 1984) and Hungary (Sziráki 1993a(Sziráki , b, 2010. Both latter authors studied the taxon carefully and arrived at the conclusion that R. mediterranea is a good species and not a subspecies of R. ophiopsis. Aside from the known and corroborated morphological differences, an important argument for the status of a separate species was the sympatry of both taxa in Romania and Hungary. The arguments of Kis (1984) and Sziráki (1993a) were convincing and accepted by us (H. Aspöck and U. Aspöck 2007, 2014. Finally, in 2013 R. mediterranea was found in the yard and on the outer walls of an old farmhouse, now representing an open-air museum, at a considerably high altitude (800 m) in Upper Austria (Rausch et al. 2016). It was an absolute surprise to find this Mediterranean snakefly in a comparatively cold region of Austria (Figs 1, 2). Moreover, R. mediterranea occurs there in an extremely high population density. It was suspected that the larvae develop within the straw covering the roof (Rausch et al. 2016), and this could recently be confirmed (Gruppe et al. 2017) (Figs 3, 4). Thus, the question arose: How has R. mediterranea achieved the establishment of a stable population in a locality in Central Europe, which offers unfavourable climatic conditions compared to many other parts of Austria (Fig. 5)? To better evaluate the phylogeographic scenarios of this species, i.e. natural expansion of the distribution range vs. human mediated dispersal, we performed molecular genetic analyses of specimens from Austria, Greece and Italy. The specimens analysed genetically were compared morphologically with specimens from many localities covering the currently known distribution. Moreover, specimens of R. ophiopsis from Upper Austria and other parts of Central Europe were included to corroborate the morphological differences between the two taxa.

Morphological studies
Numerous adults of both sexes of Raphidia mediterranea from many localities in Greece, Italy, Anatolia, as well as specimens of R. ophiopsis from Upper Austria and other parts of Central Europe were compared with imagines from Pelmberg (Upper Austria) based on the well-known morphological characters of male and female genitalia (H. Aspöck et al. 1991). Genital segments were cleared in KOH and processed in the usual manner described elsewhere.
The distribution map was provided with ArcGis/ ArcMap ver. 10.3.1.4959 based on the distribution records provided in the Suppl. material 2. Source of the map: National Geographic-Weltkarte -Content may not reflect National Geographic's current map policy. Sources: National Geographic, Esri, DeLorme, HERE, UNEP-WCMC, USGS, NASA, ESA, METI, NRCAN, GEBCO, NOAA, increment P Corp.

Molecular genetic analysis
For DNA analysis samples of four individuals of Raphidia mediterranea were selected, which had been collected in Pelmberg (Upper Austria), Gargano (Italy) and Zachlorou (Peloponnesus, Greece). Moreover, five representatives of the genus were included: Raphidia ophiopsis Linnaeus, 1758, Raphidia alcoholica H. Aspöck & U. Aspöck, 1969, Raphidia ulrikae H. Aspöck, 1964, Raphidia ariadne H. Aspöck & U. Aspöck, 1964, and Raphidia ligurica Albarda, 1891. A list of specimens analysed with exact localities is given in Table 1. Tissue samples were taken from one leg of alcohol-preserved specimens with sterile forceps. Vouchers are stored at the Entomological Department of the Museum of Natural History Vienna (NHMW). Remaining DNA is stored in the DNA and Tissue Collection of the Central Research Laboratories at the NHMW.

Marker sequences and laboratory procedures
Two mitochondrial marker sequences were amplified using primers listed in Table 2: (1) A partial sequence of the cytochrome c oxidase subunit 3 gene (cox3) which has been also used in a previous study on Neuropterida, as well as Raphidioptera (Haring and Aspöck 2004;Haring et al. 2011) and (2) the complete sequence of the cytochrome c oxidase subunit 1 gene (cox1) plus partial sequences of the adjacent tRNA genes. In addition, a partial sequence of the 28S rRNA gene (28S) was used as a nuclear marker sequence. The fragment lengths of cox1 sequences ranged from 1604 -1610 bp (due to indels in the flanking tRNA genes). The amplicon length of the cox3 sequence was 712 bp. The amplicon length of the 28S sequence was 1155-1161 bp.

Phylogenetic analyses
Raw sequences were manually aligned in BioEdit v.7.1.3 (Hall 1999) and checked for errors. The alignment was straightforward for the three marker sequences and was done in BioEdit v.7. and 28S sequences have been published in our previous paper (Haring et al. 2011;HM543275;HM543340;Agulla adnixa). The complete cox1 sequence was derived from GenBank (FJ207460.1; Agulla sp.). By comparing this sequence with published partial cox1 sequences of Agulla adnixa (e.g., KR141904.1), we deduced that the sequence FJ207460.1 is derived from Agulla adnixa (which has an identical sequence). As a result, in the concatenated data set, the outgroup sequence was derived from different individuals of the same species, which however appears to be unproblematic in this case. Bayesian Inference (BI) was used for calculating phylogenetic trees. For BI the best fitting substitution model was determined for each of the three genes as well as codon positions of the protein coding genes by jModelTest v.2.1.5 (Darriba et al. 2012) with the corrected Akaike information criterion (AICc). The BI analyses were calculated using MrBayes v.3.2.1 (Huelsenbeck and Ronquist 2001;Ronquist and Huelsenbeck 2003). Phylogenetic trees were also calculated from a combined alignment in which all three marker sequences were concatenated (length of alignment: 3345 positions). BI analyses were run for 7x10 6 generations (2 runs each with 4 chains, one of which was heated), sampling every hundredth tree. The first 25% of trees were discarded as burnin and from the remaining trees a 50% majority rule consensus tree was calculated. In addition, Neighbour Joining (NJ) trees (Saitou and Nei 1987) were calculated. Nodal support of NJ trees was evaluated with nonparametric bootstrapping based on 1000 replicates. These trees are shown to illustrate p distances among taxa in comparison of the three marker sequences.

Results
In former studies (H. Aspöck et al. 1991 and unpublished), based on male and female genitalia, populations of Raphidia mediterranea from various localities in Greece, Anatolia, and Italy could not be differentiated from each other. This was confirmed again on the basis of more material, particularly specimens from Pelmberg (Upper Austria) whose morphological characters coincide perfectly with those figured in H. Aspöck et al. (1991). Specimens of the genus Raphidia from other parts of Central Europe (except Raphidia ulrikae) proved to be conspecific with Raphidia ophiopsis.
The DNA sequence analysis revealed that the four specimens of R. mediterranea are identical in cox1 and 28S, while in cox3 a single substitution differentiating Ramed-4 from the other (identical) sequences was found. In general, the variation within 28S was extremely low. Except R. ligurica, which shows distances to the other ingroup taxa of 1.08 and 1.35%, respectively, sequences of all other ingroup species differ with p distances below dez.pensoft.net 1% or are even identical. Between Raphidia and Agulla 28S distances ranged from 6.0 to 6.5%. Concerning the mitochondrial marker sequences, p distances between R. mediterranea and R. ophiopsis (the closest relative) were 5.08% in cox1and 5.62% in cox3, while the other species of Raphidia differed between 8.63-14.83% (cox1) and 7.46-16.94% (cox3) from R. mediterranea. Distances between species in the various gene sequences are illustrated by the NJ trees in Suppl. material 1.
To assess the systematic position of R. mediterranea not only on the basis of morphological characters, we performed a phylogenetic analysis based on three genes (cox1, cox3, 28S). The two mt sequences resulted in trees in which the sister group of R. mediterranea is R. ophiopsis. In most analyses, R. alcoholica is the sister group of those two lineages, followed by R. ariadne; only in the BI tree of cox1 the relationships were unresolved (Suppl. material 1). With respect to the relationships of the other species there is a difference concerning R. ligurica and R. ulrikae depending on the marker sequence and the method applied. In some trees R. ulrikae splits from the most basal node, in others it is R. ligurica. Yet, in all trees this node is poorly supported. The tree based on 28S sequences (Suppl. material 1) is congruent with the mt based trees, yet, due to the low variation within this gene, the amount of phylogenetic information is limited. In a BI tree based on the combined marker genes (Fig. 6) all nodes are highly supported.

Discussion
The discovery of an isolated and unusually large population of Raphidia mediterranea -a Mediterranean species which has never been found elsewhere in Central Europe -in a farmhouse in a comparatively climatically unfavourable part of Upper Austria had raised the question concerning the origin of this population. It was assumed that morphological and/or genetic differences would be found, if the species had reached Upper Austria long ago by natural means of expansion of the distribution range. Therefore, specimens of the population from Upper Austria were compared with specimens from Greece and Italy. In the present study, the morphology-based results were clearly confirmed by molecular genetic analyses: The four specimens of R. mediterranea (two from Pelmberg (Austria), one from Greece, one from Italy) had almost identical sequences. It is legitimate to conclude that these populations were not separated long ago. The other species of Raphidia are clearly separated (see Fig. 6).
Substantial differences could not be found in morphological characters, particularly in male and female genitalia, or in the sequences of three genes (cox1, cox3, 28S). This implies that all presently known and examined populations of R. mediterranea originated from a single glacial refugium. This refugium can reasonably be assumed to be in the south of the Balkan Peninsula as a part of the large balkanopontomediterranean refugium (H. Aspöck et al. 1991). From there the species reached other parts of Europe (and Anatolia) not long ago. Natural dispersal from the south of the Balkan Peninsula to isolated areas of the north of the Balkan Peninsula, to southern parts of Italy, to parts of Eastern Europe and particularly parts of Central Europe is highly unlikely. Consequently, an anthropogenic introduction into various regions is highly probable. Raphidia mediterranea is a euryoecious species, whose larvae live mainly in the detritus of roots of bushes, but sometimes also under bark.
In Greece, the species occurs in many regions, in various habitats at altitudes of 10 -1200 m and often in high population densities. Thus, it might have been occasionally transported to new habitats by ships carrying wood or soil. This could have occurred already in antiquity and throughout the past centuries.
The discovery of the isolated population of R. mediterranea in Upper Austria and the failure to detect any morphological or genetic differences between these vastly distant populations supports our previous hypothesis (Aspöck et al. 1977(Aspöck et al. , 1980(Aspöck et al. , 1991(Aspöck et al. , 2001 of unintentional introduction by human activities. A natural dispersal -per continuitatem or by wind -can convincingly be excluded. How did R. mediterranea come to Upper Austria? We now know definitely that the larvae develop within the straw of the roof (Gruppe et al. 2017). Thus, it is a reasonable assumption that this snakefly was introduced with straw from somewhere on the Balkan Peninsula. The straw presently on the roof is from Austria, but in the past it may have been imported. It is also possible that live adults (theoretically one female would be sufficient) were introduced (e.g. via car, truck or bus) from the Mediterranean region to Pelmberg and subsequently the female laid eggs in the straw. Until now, no studies have been published indicating that larvae develop in straw on roofs. In Central Europe thatched roofs have become rare, but in eastern and southeastern parts of Europe such roofs are still frequent in certain regions. It would be easy and exciting to examine these habitats for snakeflies. It would particularly be interesting to know whether other species of Raphidioptera can also develop in straw of thatched roofs where they would feed on mites, spring-tails, Psocoptera, larvae of beetles and other small arthropods living in the straw.
Concerning the systematic position of R. mediterranea, the phylogenetic analysis based on three genes confirmed our view of the systematics of R. ophiopsis, R. mediterranea, R. alcoholica, R. ariadne, R. ulrikae andR. ligurica (H. Aspöck et al. 1991, 2001). It is of particular interest that R. alcoholica is the sister species of R. ophiopsis + R. mediterranea, thus confirming the close relationship of the three taxa once regarded as subspecies of R. ophiopsis.
We know little about the formation of distribution patterns of Raphidioptera. Snakeflies are generally characterised by low, in many cases extremely low, expansivity, and many species have hardly enlarged their distribution beyond their glacial refugial areas. In Central Europe, 16 species of Raphidioptera (13 Raphidiidae and 3 Inocelliidae) occur, 10 of these are of Mediterranean origin and presumably have reached Central Europe after the last glacial period, i.e. within the past 10,000 years (H. Aspöck 2008, H. Aspöck et al. 1991, H. Aspöck and U. Aspöck 2015. At least in Austria, R. mediterranea must be regarded as a human introduced neozoon.
The genera of Poliina are Holarctic (Polia) or only Eurasiatic with centre of diversity in themonsoonic mountainous areas of South-Eastern Asia (Himalayan -Sino-Tibetan faunal type). Twenty-six species of Polia are present in Eurasia and thirteen species in North America; from themonly one is a Holarctic, circumpolar species (P. richardsoni (Curtis, 1835)). Further genera of the subtribe are exclusively Palaearctic.The most diverse genus is Ctenoceratoda with more than thirty, mostly Central Asiatic species.The members of this subtribe have a characteristic "ground plan" of genital structures (including some lock-and-key mechanisms, see Varga 1992;Varga and Ronkay 2013) with several shared apomorphies as the identical structure of ampulla-harpe complex, the regularly (Polia, Haderonia) asymmetrical saccular processes covered by specialised brushes, the long, tubular endophallus (vesica) without subbasal diverticulum and cornutus but with long medio-subterminal field of fasciculate cornuti (males), the globular corpus bursae and the tubular appendix bursae (females). Abdominal brush-organs of males are mostly present and the last abdominal segment of females often shows specific strongly sclerotised, often shieldshaped structures.
In this review, based on the presence of the T-shaped vesica and the subbasal diverticulum with cornutus,typifying numerous Mamestrina genera (Varga and Ronkay 1991), but also on several characters of the genital capsula (see below), which are categorically absent in Poliina, certain genera and species formerly associated with Poliina have been excluded from this subtribe. The genus Kollariana includes three large, externally confusingly Polia-like species, the genitalia of which demonstrate, however, their close relationship with the Sideridis clade of the subtribe Mamestrina Hampson, 1902. They do not havee.g. saccular processes and ampullae on the valvae, but an ear-shaped costal process near to the cucullus. They also have claw-or spine-like sclerotisation of carina; the vesica is T-shaped, with long subbasal diverticulum and acute cornutus. Kollariana species do not have in the female genitalia elongate tubular appendix bursae, as most genera of Poliina, but they have two complete and one shorter row of small, elliptical stigmata on the corpus bursae.This genus is transferred, therefore, into Mamestrina. It is worth to mention that there are some additional large-sized, Polia-like species occurring in the mountains of the SE frontier of the Tibetan plateau (e.g. the taxa of the genus Irene, the two members of the newly described genus Multisigna ("Polia") costirufa Draudt, 1950 and "P." hofer (Saldaitis, Benedek & Behounek, 2016), and the still less investigated "Hyssia" hadulina Draudt, 1950, etc.) which all belong to Mamestrina based on the shared characters mentioned above (see: taxonomic part in details).
The male genitalia are essentially similar to those of P. (A.) mortua but the sclerotised medial extension of valvae is less falcate, evenly broad, except the finely tapering and terminally pointed distal quarter. This process is medially narrower and distally dilated, apically rounded in all four subspecies of P. (A.) mortua. In addition, the clavi are broader and more evenly rounded, and the juxta is smaller and thinner than in different subspecies of P. (A.) mortua.
Distribution. SE Tibetan. The species is known from the type-locality only; the holotype specimen was collected in a high altitude forest region in the midsummer period. It contains five easily distinguishable species representing three main lineages, the culta-, the subviolacea-and the kalikotei-lineages. The shortened diagnosis of the subgenus is presented below; the detailed analysis of the clade is given in the above-mentioned publication. Diagnosis. Metallopolia species are large, robust moths, resembling the larger south Siberian Polia species but have shorter abdomen bearing 3-4 prominent blackish tufts on the first abdominal segments dorsally. The most conspicuous external character of the members of this subgenus is the presence of optically structured "metallic" scales with "neon-greenish" colouration (see: Etymology) within or near to the maculation and the anal edge of the postmedial transversal line. The forewing ground colour is rather dark brown to blackish-brown with some purplish-violaceous hue and diffuse, smaller or larger reddish-brownish patches; the hindwings are also dark brown or grey-brown. The members of the subgenus are externally often confusingly similar, the proper identification often requires the study of genitalia.

Subgenus
In the male genitalia, the saccular processes are slightly asymmetrical, extended, acute or obtuse, with strong setae terminally, in most species with characteristic brush of specialised setae on the right (in figures left) side. Vesica long, tubular, partly or entirely coiling, medial and distal sections armed by numerous small, spiniform cornuti arranged into a long and variably dense stripe. In the female genitalia, the ductus bursae is sclerotised, compressed dorso-ventrally; the appendix bursae is tubular, sausage-shaped, slightly retroflexed, bursa globular with longitudinal, extremely faint signa.
Etymology. The name refers to the scales with light greenish optical colouration and metallic shine on the fore wings as unique character within the genus Polia. elongate triangular and somewhat acute forewings, longer pectinated and relatively long antennae of males, by dark greyish-brown colouration of thorax and fore wings with some violaceous shine.
The male genitalia also differ conspicuously from those of all other known Polia species, the diagnostic features are as follows: the saccular processes are symmetrical, relatively short and densely covered by a "bush" of strong setae terminally, and the vesica is completely helicoid and recurved, bearing a large number of thin, spiniform cornuti and a small terminal diverticulum.
The female genitalia are also very specific: the sclerotisation of the antrum is weak, connected to ductus bursae with a slightly constricted membranous section ("neck"), the ductus bursae is flattened and more sclerotised, the corpus bursae is saccate, lacking signa, while the appendix bursae is broad and retroflexed.

Pachetra sagittigera (Hufnagel, 1766)
Phalaena sagittigera Hufnagel, 1766, Berlinisches Magazin 3 (3) Taxonomic notes. P. cherrug shows in the genitalia of both sexes a very close relationship with P. sagittigera. The shared characters are as follows: in male genitalia the similar and unusual shape of the cucullus, the very long tubular and completely helicoid vesica nearly completely covered by a long and broad stripe of dense spinulose structures; in female genitalia the long and broad, dorso-ventrally flattened ductus bursae, the similar shape of the appendix bursae and one long and one shorter stripe of sigma. Both species are also bionomically closely related, they have grass-feeding larvae in contrast to the dicot herbaceous and woody food plants of Polia spp. The distribution and certain taxonomic questions are discussed in detail by Dinca (2010).

Genus Tricheurois Hampson, 1905
Tricheurois Hampson, 1905 (Draudt, 1950).  Taxonomic notes. Despite of the "Polia-like" habitus the genital structures of both sexes clearly show that Kollariana belong to the "Sideridis" line of the subtribe Mamestrina (See also: Introduction). In the male genitalia, the genital capsule is very similar to that of certain large Sideridis species as e.g. S. turbida (Esper, 1790) or S. egena (Lederer, 1853). The diagnostic features are as follows: valva without saccular process and ampulla, but with ear-shaped costal process near to cucullus; aedeagus with clawor spine-shaped sclerotisation of carina; vesica T-shaped, with long subbasal diverticulum and acute cornutus. In the female genitalia there are two complete and one shorter row of small, elliptical stigmata on the corpus bursae; the ductus bursae is rather strongly sclerotised with lateral pouch corresponding to the sclerotised extension of carina.
Polia albomixta Draudt, 1950 fig. 9, here designated. Diagnosis. The two species of this new genus show some external similarity with the larger Polia species (as e.g. P. vesperugo) but they also resemble the larger Sideridis species (as e.g. S. egena, S. turbida) but also Apamea species according the robust body, dark brownish-greyish colouration and pattern of fore wing with regular maculation and crenulate or zigzag-shaped crosslines. The most important differential characters are in the genitalia of both sexes. In male genitalia the saccular part of valvae is not extended, no any trend of asymmetrisation and/or presence of specialised saccular brushes can be observed. Free "clasper" of the harpe-ampulla complex -which is usually present in Polia -is reduced. Vesica is not elongate-tubular as in Polia, but it shows the modified form of the T-shaped vesica of the Sideridis-clade of the Mamestrina while it is more saccate with subbasal diverticulum and the subterminal field of cornuti is transformed to a single huge cornutus, unusual for other related genera. The female genitalia are also strikingly different from Polia but also from the genera of the subtribe Mamestrina (e.g. Sideridis, Conisania which seem to be most closely related) by the presence of very numerous pearl-shaped signa and by the conical appendages of ductus bursae. Based on these characters we place this genus near to the also habitually similar genera Kollariana and Irene.
Etymology.The generic name refers to the most peculiar character of the female genitalia. ( (Guenée, 1852) was associated with Lacanobia and S. grandis (Guenée, 1852) was introduced to this genus (Lacanobia grandis).

Multisigna hofer
Tycomarptes proximoides (Wiltshire, 1982) Haderonia proximoides Wiltshire, 1982 This year we look back at 160 years of entomological research published in the DEZ. Believe it or not, our journal is the third oldest of all still existing entomological periodicals worldwide! A concatenation of favourable circumstances? At first glance, the first decades were rather tough ones, involving personal controversies, splitting of the society behind the journal and the journal itself, and later reunion (Wessel 2007). However, at the second glance, this period seems to have been an excellent one at the same time, as the young and dedicated visionary, Gustav Kraatz, the first editor of the DEZ, guided the journal throughout these troublesome waters for the first 50 years. What makes him visionary? Already 160 years ago, he promoted high standards in taxonomical publications such as the description of both sexes when erecting new genera, the publication of comprehensive revisions instead of single species descriptions and the exploration of new diagnostic characters (Wessel 2007) -not much to add 160 years later! More than this, under his editorship the DEZ was at the forefront of the development and establishing of internationally recognized nomenclatorial rules in entomology, regulating foremost issues of priority (Wessel 2007). Still today, nomenclatorial issues constitute a hot topic in entomological publishing. Finally, Gustav Kraatz was driven by the urge to combine collections and libraries of all German entomologists, so that scientists could have free access (Wessel 2007). Together let's do our best to continue this line to the future!