Research Article
Print
Research Article
The Nevrorthidae, mistaken at all times: phylogeny and review of present knowledge (Holometabola, Neuropterida, Neuroptera)
expand article infoUlrike Aspöck§, Horst Aspöck|, Xingyue Liu
‡ Natural History Museum Vienna, Vienna, Austria
§ University of Vienna, Vienna, Austria
| Medical University (MUW), Vienna, Austria
¶ China Agricultural University, Beijing, China
Open Access

Abstract

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.

Sonnet for a Vulnerable Creature

Is the Climate still fine?

Still clean, the Riverine?

Ruined rivulets run dry

Fossils – tho’ living – may Die.

Key Words

systematics, biology, distribution, biogeography

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” (1906–1908), Nevrorthidae were still cryptic and hidden within Sisyridae (as Sisyra (Rophalis) relicta Hagen, 1856).

Figure 1. 

Nevrorthus iridipennis Costa. Italy: Calabria. From Costa (1863), who published the first illustration of a nevrorthid.

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 1992, 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 compact 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. (2013, 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.

Figure 2. 

Photographs of living nevrorthids. a Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel, female, Italy: Friuli (Photo P. Sehnal) b Nevrorthus apatelios, larva, Italy: Friuli (Photo F. Denner (former Anderle) c Nipponeurorthus fuscinervis (Nakahara), female, Japan, Hokkaido (Photo X. Liu) d Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, female holotype, China, Yunnan (Photo H. Li).

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, 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 (1845-1856), Nel and Jarzembowski (1997), Makarkin and Perkovsky (2009), Wichard et al. (2009, 2010), Wedmann et al. (2013), Wichard (2014, 2016), Makarkin (2016).

Material and methods

List of taxa examined

Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel, 1977

Nevrorthus fallax (Rambur, 1842)

Nevrorthus hannibal U. Aspöck & H. Aspöck, 1983

Nevrorthus iridipennis Costa, 1863

Austroneurorthus brunneipennis (Esben-Petersen, 1929)

Austroneurorthus horstaspoecki U. Aspöck, 2004

Nipponeurorthus damingshanicus Liu, U. Aspöck & U. Aspöck

Nipponeurorthus fasciatus Nakahara, 1958

Nipponeurorthus flinti U. Aspöck & H. Aspöck, 2008

Nipponeurorthus furcatus Liu, H. Aspöck & U. Aspöck, 2014

Nipponeurorthus fuscinervis (Nakahara, 1915)

Nipponeurorthus multilineatus Nakahara, 1966

Nipponeurorthus pallidinervis Nakahara, 1958

Nipponeurorthus punctatus (Nakahara, 1915)

Nipponeurorthus tinctipennis Nakahara, 1958

Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, 2012

Laboratory techniques

Photographs of living adults were made with a Nikon D300 or D90 with a Nikon AF Micro-NIKKOR 105mm f/2.8D lens and Nikon macro flash -Kit R1 (Figs 1a, c, d). 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.

Figure 3. 

Wings and genital segments of representatives of the genera Nevrorthus Costa and Austroneurorthus Nakahara. a Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel, male paratype, Greece: Peloponnesus/Peloponnese (Photo H. Bruckner) b Nevrorthus fallax (Rambur), female, Italy: Sardinia (Photo H. Bruckner) c Nevrorthus hannibal U. Aspöck & H. Aspöck, male holotype, Tunisia: S Ain Draham (Photo H. Bruckner) d Nevrorthus iridipennis Costa, male, Italy: Sicilia (Photo H. Bruckner) e–f Nevrorthus reconditus Monserrat & Gavira e right fore- and hindwing f male genital segments, ventral, Spain: Malaga (adapted from Monserrat and Gavira 2014) g Austroneurorthus brunneipennis (Esben-Petersen), male paratype, Australia: Tambourine Mt. (Photo H. Bruckner) h Austroneurorthus horstaspoecki U. Aspöck, male, Australia: Victoria (Photo H. Bruckner). Abbreviations. A– Analis; C – Costa; CuA – Cubitus anterior; CuP – Cubitus posterior; MA – Media anterior; MP – Media posterior; R – Radius; Rs – Radial sector; Sc – Subcosta. Scale bar: 1.0 mm.

Figure 4. 

Wings of representatives of the genera Nipponeurorthus Nakahara and Sinoneurorthus Liu, H. Aspöck & U. Aspöck. a. Nipponeurorthus fasciatus Nakahara, male, Taiwan: Nantou (Photo H. Bruckner); b. Nipponeurorthus fuscinervis (Nakahara), Japan: Aomori (Photo X. Liu); c. Nipponeurorthus damingshanicus Liu, H. Aspöck & U. Aspöck, female paratype, China: Guangxi (Photo X. Liu); d. Nipponeurorthus furcatus Liu, H. Aspöck & U. Aspöck, male paratype, China: Yunnan (Photo X. Liu); e. Nipponeurorthus flinti U. Aspöck & H. Aspöck, male, Japan: Amamioshima (Photo X. Liu); f–g. Nipponeurorthus pallidinervis Nakahara, f: male paratype, g: female paratype, Japan: Hokkaido (Photo H. Bruckner); h. Nipponeurorthus multilineatus Nakahara, male, Taiwan (Photo H. Bruckner); i. Nipponeurorthus punctatus (Nakahara), male, Japan (Photo X. Liu); j. Nipponeurorthus tinctipennis Nakahara, male, Japan: Yakushima Island (Photo X. Liu); k. Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, female holotype, China: Yunnan (Photo X. Liu). Abbreviations. A – Analis; C – Costa; CuA – Cubitus anterior; CuP – Cubitus posterior; MA – Media anterior; MP – Media posterior; R – Radius; Rs – Radial sector; Sc – Subcosta. Scale bars: 1.0 mm.

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-WCMC, 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.

Results

Taxonomy

List of abbreviations

anat (anatomy), annotcat (annotated catalogue), biogeogr (biogeography), biol (biology), cat (catalogue), charact (characteristics), com (comment), compmorphol (comparative morphology), descr (description), distr (distribution), distrmap (distribution map or maps), ecol (ecology), fig (figure), gs (genital segments), key (identification key), la (larvae), list (listed or mentioned), mon (monograph), nom (nomenclature), odescr (original description), overv (overview), phyl (phylogeny), pu (pupae), rec (record), syn (synonymisation), syst (systematics), tax (taxonomy).

Acronyms of institutions

Entomological Museum, China Agricultural University, Beijing, China (CAU); National Science Museum, Tokyo, Japan (NSMT); Australian Museum, Sydney, Australia (AMS); Australian National Insect Collection, Canberra, Australia (CSIRO); Smithsonian Institution, National Museum of Natural History, Washington D.C., USA (NMNH); Texas A & M University, College Station, Texas, USA (TAM); Zoologisk Museum, Copenhagen, Denmark (ZMC); Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt-Universität zu Berlin, Berlin, Germany (MFN); Naturhistorisches Museum Wien, Vienna, Austria (NHMW); Museo Zoologico dell’Universitá di Napoli Federico II, Naples, Italy (MZUN); Collection of Horst & Ulrike Aspöck, Vienna, Austria (HUAC); Collection of Victor Monserrat, Madrid, Spain (VM); Collection of Fumio Hayashi, Tokyo, Japan (HFC).

Nevrorthidae Nakahara, 1915

Neurorthini Nakahara, 1915: 14 (nom).

Neurorthinae Nakahara: Nakahara 1958 (mon, nom).

Neurorthidae Nakahara: Zwick 1967 (la, compmorphol, syst); Gaumont 1968 (compmorphol la); Riek 1970(charact); Gaumont 1976 (compmorphol la); Monserrat 1977 (nom, list); H. Aspöck et al. 1978 (charact); New 1978 (ecol la); H. Aspöck et al. 1980 (mon); Henry 1982 (charact); Gepp 1984 (tax la); Malicky 1984 (biol, ecol); New 1986 (charact, biol), 1989 (tax), 1991 (charact, tax, tax la); U. Aspöck 1995 (phyl); Güsten 1996 (compmorphol); New 1996 (cat: Australia); Wachmann and Saure 1997 (tax, tax la).

Nevrorthidae Nakahara: Oswald and Penny 1991 (list, nom); U. Aspöck 1995 (phyl); H. Aspöck and Hölzel 1996 (overv); U. Aspöck and H. Aspöck 1999 (overv); H. Aspöck et al. 2001 (annotcat); U. Aspöck et al. 2001 (phyl); U. Aspöck and H. Aspöck 2007 (biogeogr, distrmap, figs: gs males); Monserrat and Gavira 2014 (distrmap).

Remarks

The Nevrorthidae are a species-poor relic family with an extremely vicariant distribution pattern (Fig. 17) of its four extant and five extinct genera: Nevrorthus Costa, 1863, comprising five disjunctively scattered Mediterranean species: N. iridipennis Costa, 1863 (Italy: Calabria, Sicily), N. apatelios H. Aspöck, U. Aspöck & Hölzel, 1977 (Balkan Peninsula, Romania, northern Italy: Friuli, and Slovenia), N. fallax (Rambur, 1842) (France: Corsica, Italy: Sardinia), N. hannibal U. Aspöck & H. Aspöck, 1983 (Algeria, Tunisia), N. reconditus Monserrat & Gavira, 2014 (Spain: Malaga); Austroneurorthus Nakahara, 1958, comprising two species, restricted to southeastern parts of Australia: A. brunneipennis (Esben-Petersen, 1929) (southeastern Queensland, New South Wales), A. horstaspoecki (U. Aspöck, 2004) (Victoria, New South Wales); Nipponeurorthus Nakahara, 1958, comprising eleven species, distributed in China and Japan: Ni. damingshanicus Liu, H. Aspöck & U. Aspöck, 2014 (China: Guangxi), Ni. fasciatus Nakahara, 1958 (China: Taiwan), Ni. flinti U. Aspöck & H. Aspöck, 2008 (Japan: Okinawa, Amamioshima), Ni. furcatus Liu, H. Aspöck & U. Aspöck, 2014 (China: Yunnan), Ni. fuscinervis (Nakahara, 1915) (Japan: Hokkaido, Honshu), Ni. multilineatus Nakahara, 1966, (China: Taiwan), Ni. pallidinervis Nakahara, 1958 (Japan: Hokkaido, Honshu, Kyushu, Tsushima Island), Ni. punctatus (Nakahara, 1915) (Japan: Honshu, Hokkaido, Kyushu, Yakushima), Ni. qinicus Yang in Chen, 1998 (China: Shaanxi), Ni. tianmushanus Yang & Gao, 2001 (China: Zhejiang), Ni. tinctipennis Nakahara, 1958 (Japan: Yakushima); Sinoneurorthus Liu, H. Aspöck & U. Aspöck, 2012, so far comprising only one described species, S. yunnanicus Liu, H. Aspöck & U. Aspöck, 2012 (China: Yunnan).

Extinct taxa from the Eocene Baltic amber are assigned to the monotypic genus Rophalis Hagen, 1856, with R. relicta (Hagen in Berendt, 1845-1856), Electroneurorthus Wichard, Buder & Caruso, 2010, comprising E. malickyi Wichard, Buder & Caruso, 2010, Palaeoneurorthus Wichard, 2009, comprising P. bifurcatus Wichard, 2009, P. hoffeinsorum Wichard, 2009, P. groehni Wichard, Buder & Caruso, 2010, P. eocaenus Wichard, 2016, Balticoneurorthus Wichard, 2016, with B. elegans Wichard, 2016, and Proberotha Krüger, 1923, comprising Pr. prisca Krüger, 1923, and Pr. dichotoma Wichard, 2016.

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.

Nevrorthus Costa, 1863

Nevrorthus Costa, 1863: 32 (odescr) [Type species: Mucropalpus fallax Rambur, 1842, by subsequent designation]:

Leraut 1981 (nom); Oswald and Penny 1991 (nom); H. Aspöck et al. 2001 (annotcat); U. Aspöck and H. Aspöck 2007 (biogeogr, distrmap); U. Aspöck and H. Aspöck 2008a (compmorphol, figs: gs female); U. Aspöck and H. Aspöck 2010a (overview, biogeogr, fig: distrmap); Gavira et al. 2012 (com); Monserrat and Trivino 2013 (com); Monserrat and Gavira 2014 (distrmap).

Neurorthus Costa (unjustified emendation): McLachlan 1881 (nom); Nakahara 1915 (charact); Klapálek 1917 (descr); Navás 1935 (mon); Parfin and Gurney 1956 (mon); Nakahara 1958 (charact); Zwick 1967 (fig: la, compmorphol, syst); Tjeder 1979 (compmorphol); H. Aspöck et al. 1980 (mon); Malicky 1984 (biol, ecol, distr); Oswald and Penny 1991 (nom); Wichard et al. 1995 (fig pu); Wachmann and Saure 1997 (key).

Sartena Hagen, 1864: 41 (odescr) [Type species: Sartena amaena Hagen, 1864, by monotypy]: McLachlan 1881 (nom); H. Aspöck et al. 1980 (syn); Oswald and Penny 1991 (nom).

Diagnosis

Adults of small body size; male forewing length 6–8 mm. Body coloration greyish-brownish. Forewings transparent to pale yellowish, crossveins sometimes dark and shaded. Costal cross veins of forewings not forked. Hindwing MA and anterior branch of MP forked distal to outer series of gradate cross veins. Male: Abdominal segment 7 enlarged. A ring-like zone of glands present between male abdominal segments 7 and 8. Abdominal eversible sacks present between segments 6 and 7. Male sternite 9 long, strongly extending posteriad; gonocoxites 9 as huge plates with digitiform gonostyli 9 and processus-like gonapophyses; complex of gonocoxites + gonostyli + gonapophyses 10 amalgamated with sternite 9, forming a pseudoapex of the latter and framing it laterally; gonocoxites 11 fused into a bow-like bridge. Female: Fused gonocoxites 8 forming a broad trapezoid sclerite; gonocoxites 9 club-shaped, without distinct gonostyli; bursa copulatrix comprising a sclerotized structure.

Distribution

Mediterranean region.

Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel, 1977

Figs 2a–b; 3a; 5a–e; 6c; 14

Neurorthus iridipennis auct. nec Costa (misidentification): Klapálek 1917 (syst, distr); Pongracz 1923 (distr); Zelený 1964 (rec); 1971 (rec).

Neurorthus apatelios H. Aspöck, U. Aspöck & Hölzel, 1977: 54 (odescr, figs: gs male): H. Aspöck et al. 1978 (distr); H. Aspöck et al. 1980 (mon); U. Aspöck and H. Aspöck 1983 (distr); Malicky 1984 (ecol, distr); Saure 1989 (distr); Popov 1990 (distr); 1991 (rec); 1992 (distr); Devetak 1992 (distr); Popov 1993 (distr);

Nevrorthus apatelios: H. Aspöck and Hölzel 1996 (distr); U. Aspöck and H. Aspöck 1999 (fig); H. Aspöck et al. 2001 (annotcat); Devetak and Jakšič 2003 (distr); Letardi et al. 2006 (distr, biol); U. Aspöck and H. Aspöck 2007 (figs: adult, distrmap); Popov 2007 (distr, biol); U. Aspöck and H. Aspöck 2008a (compmorphol, fig: gs female); Sziráki 2008 (rec, distr); Jones and Devetak 2009 (distr); U. Aspöck and H. Aspöck 2010a (biogeogr, fig: distrmap); Monserrat and Gavira 2014 (figs: gs, head, thorax); Devetak and Klokočovnik 2016 (biol).

Type locality

Greece (Euboea: S Prokopion).

Male

Body length 2.2 mm; forewing length 6.0–7.5 mm, hindwing length 5.5–6.5 mm.

Head yellowish. Antennae pale yellow, scapus and pedicellus brownish. Mouthparts yellow.

Prothorax yellow; meso- and metathorax darker. Legs yellow. Wings hyaline, membrane uncoloured; forewing veins yellowish; hindwing veins pale yellow, paler than in forewing.

Abdomen dorsally dark brown with yellow pattern, ventrally yellowish. 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 rounded. Gonocoxites 11 fused into a bow-like bridge.

Female

Body length 2.4 mm; forewing length 7.4–7.9 mm, hindwing length 7.6–7.8 mm.

Fused gonocoxites 8 forming a broad trapezoid sclerite; fused gonapophyses 8 triangular; gonocoxites 9 club-shaped, without distinct gonostyli; bursa copulatrix comprising a sclerotized structure.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): “Griechenland, Euboea, S von Prokopion, 38°42’N / 23°30’E, 250 m, 24.5.1974, H. Malicky leg.” (HUAC).

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).

Distribution

Albania, Bosnia-Herzegovina, Bulgaria, Greece, Italy, Kosovo, Macedonia, Romania, Serbia, Slovenia.

Figure 5. 

Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel. a. Male, genital segments, lateral; b. Same, ventral; c. Female, genital segments, lateral; d. Gonocoxites 8 and gonapophyses 8, ventral; e. Bursa copulatrix, lateral. Scale bars: 0.5 mm.

Figure 6. 

Male genital segments of Nevrorthus spp. a Nevrorthus fallax (Rambur), ventral b Nevrorthus iridipennis Costa, ventral c Nevrorthus apatelios H. Aspöck, U. Aspöck & Hölzel, ventral d–e Nevrorthus hannibal U. Aspöck & H. Aspöck, male holotype: d lateral e ventral f–g Nevrorthus fallax (Rambur), female: f gonocoxites 8 and gonapophyses 8, ventral g lateral h–i Nevrorthus hannibal U. Aspöck & H. Aspöck, female: h gonocoxites 8 and gonapophyses 8, ventral i lateral. Abbreviations. e – ectoproct; g – ring of glands; gp – gonapophysis; gs – gonostylus; gx – gonocoxite; p – pleuritocava; S – sternite; T – tergite. Scale bars: 0.5 mm.

Nevrorthus fallax (Rambur, 1842)

Figs 3b; 6a, f–g; 14

Mucropalpus fallax Rambur, 1842: 422 (odescr).

Sartena amaena Hagen, 1864: 41 (odescr). McLachlan 1881 (nom).

Neurorthus fallax (Rambur): McLachlan 1881 (nom); 1898 (com); Klapálek 1917 (descr, distr, figs: wings, gs male); Esben-Petersen 1913 (distr); Lestage 1924 (syst); Kimmins 1930 (distr); Nakahara 1958 (charact, figs wings, gs male, female); Zwick 1967 (ecol, compmorphol, syst, figs: gs male, la); Gaumont 1976 (anat); H. Aspöck et al. 1977 (tax); Tjeder 1979 (tax); H. Aspöck et al. 1980 (mon); U. Aspöck and H. Aspöck 1983 (distr); Malicky 1984 (ecol, distr, tax, figs: la); Letardi 1994 (distr).

Nevrorthus fallax (Rambur): Leraut 1981 (distr); Pantaleoni 1994 (distr); Iori et al. 1995 (distr); H. Aspöck and Hölzel 1996 (distr); U. Aspöck and H. Aspöck 1999 (fig: la); H. Aspöck et al. 2001 (annotcat); U. Aspöck and H. Aspöck 2007 (fig: distrmap); Letardi et al. 2008 (rec); U. Aspöck and H. Aspöck 2010a (biogeogr, fig: distrmap); Monserrat and Gavira 2014 (figs: gs, adult).

Type locality

Italy (Sardinia).

Male

Body length 2.2 mm; forewing length 6.0–8.0 mm, hindwing length 5.5–6.5 mm.

Head yellowish, dark brown line at middle. Antennae pale yellow, scapus and pedicellus brownish. Mouthparts yellow.

Prothorax yellow; meso- and metathorax darker. Legs yellow. Wings hyaline, membrane uncoloured; forewing veins yellowish; hindwing veins pale yellow, paler than in forewing.

Abdomen dorsally dark brown with yellow pattern, ventrally brown. 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 sinuated. Gonocoxites 11 fused into a bow-like bridge.

Female

Body length 2.4 mm; forewing length 7.4–7.9 mm, hindwing length 7.6–7.8 mm.

Fused gonocoxites 8 forming a broad trapezoid sclerite; fused gonapophyses 8 triangular; gonocoxites 9 club-shaped, without distinct gonostyli; bursa copulatrix comprising a sclerotized structure.

Specimens examined and records published

Supplementary material 1. Original type(s) lost. Herewith, a male is designated as neotype: “6.6.–11.6. 1978, Sardinien (I) Monti di Gennargentu 700m 40°06’N / 9°32’E, H.u.L. Hölzel leg.” (NHMW).

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) reports findings of larvae in waters with temperature 10.7–20.1°C (Sardinia) and 8.6–21.4°C (Corsica).

Distribution

Italy (Sardinia), France (Corsica).

Nevrorthus hannibal U. Aspöck & H. Aspöck, 1983

Figs 3c; 6d–e; 6h–i; 14

Neurorthus fallax McLachlan nec Rambur (misidentification: McLachlan 1898 (com))

Neurorthus iridipennis Klapálek nec Costa: Klapálek 1917 (com).

Neurorthus hannibal U. Aspöck & H. Aspöck, 1983 (odescr, figs: gs male, female, distrmap); Malicky 1984 (rec, ecol).

Nevrorthus hannibal: H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (annotcat); U. Aspöck and H. Aspöck 2007 (fig: distrmap); U. Aspöck and H. Aspöck 2010a (biogeogr, fig: distrmap); Monserrat and Gavira 2014 (figs: gs, forewing, head and thorax).

Type locality

Tunisia (S Ain Draham).

Male

Body length 2.2 mm; forewing length 7.0–8.0 mm, hindwing length 5.5–6.5 mm.

Head yellowish, vertex caudally darker. Antennae pale yellow, scapus and pedicellus brownish. Mouthparts yellow.

Prothorax yellow; meso- and metathorax darker. Legs yellow. Wings hyaline, membrane uncoloured; forewing veins yellowish; hindwing, veins pale yellow, paler than in forewing.

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.

Female

Body length 2.4 mm; forewing length 7.2–8.2 mm, hindwing length 7.6–7.8 mm.

Fused gonocoxites 8 forming a broad trapezoid sclerite; fused gonapophyses 8 triangular; gonocoxites 9 club-shaped, without distinct gonostyli; bursa copulatrix comprising a sclerotized structure.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): “Tunesien, 4 km S Ain Draham, 36°43’N / 8°40’E, 530 m, 17.–18.V.1982 (T6), H. Malicky leg.” (HUAC).

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).

Distribution

Tunisia, Algeria.

Nevrorthus iridipennis Costa, 1863

Figs 1; 3d; 6b; 14

Nevrorthus iridipennis Costa, 1863: 33 (odescr, fig: wings); Iori et al. 1995 (distr); H. Aspöck and Hölzel 1996 (distr); Letardi and Pantaleoni 1996 (distr); Pantaleoni 1999 (lectotype); H. Aspöck et al. 2001 (annotcat); Pantaleoni 2005 (com); U. Aspöck and H. Aspöck 2007 (fig: distrmap); U. Aspöck and H. Aspöck 2010a (fig: distrmap); Nicoli Aldini et al. 2012 (com); Monserrat and Gavira 2014 (figs: gs, head, thorax).

Neurorthus iridipennis Costa: McLachlan 1881 (nom); 1898 (com); Klapálek 1917 (descr, distr, figs: gs male, female); Lestage 1924 (sys); Nakahara 1958 (com); Principi 1966 (distr, rec); H. Aspöck et al. 1977 (tax); H. Aspöck et al. 1980 (mon); U. Aspöck and H. Aspöck 1983 (distr); Malicky 1984 (ecol, biol, distr, rec; figs: cocoon, pu); Monserrat 1985 (nom); Letardi 1994 (distr); Wichard et al. 1995 (fig: pu); Nicoli Aldini et al. 2012 (rec).

Type locality

Italy (Calabria).

Male

Body length 2.25 mm; forewing length 6.5 mm, hindwing length 5.5–6.5 mm.

Head yellowish, vertex caudally darker. Antennae pale yellow, scapus and pedicellus brownish. Mouthparts yellow.

Prothorax yellow; meso- and metathorax darker. Legs yellow. Wings hyaline, membrane uncoloured; forewing veins yellowish; hindwing veins pale yellow, paler than in forewing.

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 deeply forked. Gonocoxites 11 fused into a bow-like bridge.

Female

Body length 2.4 mm; forewing length 7.2–8.2 mm, hindwing length 7.6–7.8 mm.

Fused gonocoxites 8 forming a broad trapezoid sclerite; fused gonapophyses 8 triangular; gonocoxites 9 club-shaped, without distinct gonostyli; bursa copulatrix comprising a sclerotized structure.

Specimens examined and records published

Supplementary material 1. Lectotype female (by explicit designation): Calabria, Reggio Calabria “Valli di Aspromonte” (MZUN), Pantaleoni (designated 1993, 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.

Distribution

Italy (Calabria, Sicily).

Nevrorthus reconditus Monserrat & Gavira, 2014

Figs 3e–f; 14

Nevrorthus reconditus Monserrat & Gavira, 2014: 352 (odescr, figs: wings, gs male, la, distrmap).

Type locality

Spain (Malaga: Coín, Sierra Alpujata).

Male

Forewing length 6.1 mm, hindwing length 5.1 mm.

Head very pale brown. Antennae pale yellow, scapus and pedicellus brownish, basal two thirds of flagellum pale brownish, apically darker. Mouthparts brownish.

Pronotum pale brownish, with irregular darker pattern; meso-metanotum pale brownish with dark brown patches. Legs brownish. Wings hyaline, membrane uncoloured; forewing veins brownish, crossveins very dark and with dark shadows; hindwing veins brownish, crossveins partly with shadow.

Abdomen with tergites and sternites irregularly brownish pigmented. 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 sinuate. Gonocoxites 11 fused into a bow-like bridge.

Female

Forewing length 6.4–6.7 mm, hindwing length 5.8–6.0 mm.

Text adapted from Monserrat and Gavira (2014): Tergite 9 with a small circular emargination on the caudal margin. Fused gonocoxites 8 forming a broad sclerite with external margins straight; gonocoxites 9 narrow and digitiform.

Specimens examined by Monserrat and Gavira (2014), see there and Supplementary material 1. Holotype male (by original designation): “Spain, Malaga, Coín, Sierra Alpujata, Arroyo del Manzano, 30SUF35 (WGS84), 450 m, 13.V.2013, captured with a light trap in perennial stream covered by bushy willow gallery forest, T. Herrera, P. Carrasco & O. Gavira leg.” (VM).

Biology and ecology

Adults have been taken from April–May. The known vertical distribution is 150–450 m. The larva is known and has been described (Monserrat and Gavira 2014).

Distribution

Spain (Malaga).

Austroneurorthus Nakahara, 1958

Austroneurorthus Nakahara, 1958: 29 (odescr) [Type species: Neurorthus brunneipennis Esben-Petersen, 1929, by original designation].

Austroneurorthus Nakahara, 1958: New 1978 (com); U. Aspöck 2004 (distr); U. Aspöck and H. Aspöck 2007 (fig: distrmap); U. Aspöck and H. Aspöck 2010a (fig: distrmap).

Diagnosis

Adults of small body size; male forewing length 6.0–8.0 mm, hindwing length 6.0–7.0 mm, female forewing length 7.8–9.0 mm, hindwing length 6.8–8.0 mm. Body coloration yellowish, with dark pattern or brownish. Forewings transparent, crossveins partly dark and shaded. Costal crossveins of forewings partly forked. Hindwing MA and anterior branch of MP forked proximal to outer series of gradate crossveins. Male abdominal segment 7 not enlarged. A ring-like zone of glands present between male abdominal segments 8 and 9. Abdominal eversible sacks absent. Male sternite 9 long, strongly extending posteriad; gonocoxites 9 as huge plates without articulated gonostyli; gonapophyses 9 forming lobes; complex of gonocoxites + gonostyli + gonapophyses 10 amalgamated with sternite 9, forming a pseudoapex of the latter and framing it laterally; gonocoxites 11 fused into a broad sclerite. Fused female gonocoxites 8 forming a rectangular sclerite; gonocoxites 9 club-shaped, without distinct gonostyli; bursa copulatrix comprising a sclerotized structure.

Distribution

Australia.

Austroneurorthus brunneipennis (Esben-Petersen, 1929)

Figs 3g; 7a–c; 15

Neurorthus brunneipennis Esben-Petersen, 1929: 33 (odescr, fig: wings).

Austroneurorthus brunneipennis (Esben-Petersen, 1929): Nakahara 1958 (nom, charact, figs: wings, gs male, female); U. Aspöck and H. Aspöck 2007 (Fig: distrmap).

Type locality

Australia (Queensland: Tamborine Mt.).

Male

Forewing length 7.0–8.0 mm, hindwing length 6.0–7.0 mm.

Head yellowish. Antennae and mouthparts yellowish.

Pronotum yellowish; meso-metanotum ochre. Legs yellowish. Wing membrane hyaline, in the original description it is characterised as “yellowish tinged; but the apical margin narrowly brownish shaded” (available material was, however, rather faded); forewing longitudinal veins brownish yellow, crossveins brownish, slightly shaded; hindwing paler than forewing, veins pale yellow.

Abdomen dorsally dark brown with yellow pattern, ventrally yellowish. Male: Gonocoxites 9 as huge plates, apically rounded, gonostyli 9 not discernible, gonapophyses 9 processus-like; ectoproct broadly rounded. Complex of gonocoxites + gonostyli + gonapophyses 10 partly amalgamated with sternite 9, forming i) a pseudoapex of the latter which is deeply forked and ii) a paired hook. Gonocoxites 11 fused into a broad plate with a big median tooth.

Female

Forewing length 8.5 mm, hindwing length 8 mm.

Fused gonocoxites 8 forming a broad trapezoid sclerite; gonocoxites 9 club-shaped, without distinct gonostyli.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): Australia: Queensland, “Tambourine Mt. 5/11/1928 (L. Franzen leg.)” (CSIRO).

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).

Distribution

Australia (NSW, Queensland).

Figure 7. 

Genital segments of Austroneurorthus spp. a–c Austroneurorthus brunneipennis (Esben-Petersen), male paratype: a lateral b ventral c caudal d–g Austroneurorthus horstaspoecki U. Aspöck, male holotype, genital segments: d lateral e dorsal f caudal g ventral h–i Austroneurorthus horstaspoecki U. Aspöck, female paratype, genital segments: h lateral i ventral. Abbreviations. b – bursa copulatrix; e – ectoproct; g – ring of glands; gp – gonapophysis; gs – gonostylus; gx – gonocoxite; S – sternite; T – tergite. Scale bars: 0.5 mm.

Austroneurorthus horstaspoecki U. Aspöck, 2004

Figs 3h; 7d–i; 15

Austroneurorthus horstaspoecki U. Aspöck, 2004: 177 (odescr); U. Aspöck and H. Aspöck 2008a (compmorphol, figs: gs male).

Type locality

Australia (Victoria: Aucheron R.).

Male

Forewing length 6.5–7.0 mm, hindwing length 5.5–6.0 mm.

Head yellowish. Antennae and mouthparts yellowish.

Pronotum yellowish; meso-metanotum ochre. Legs yellowish, femora on inner side with dark ovoid plate with smooth surface. Wing membrane hyaline, slightly smoky; forewing longitudinal veins yellowish, crossveins brownish, partly “shaded”. Hindwing paler than forewing, crossveins brownish.

Abdomen dorsally dark brown with yellow pattern, ventrally yellowish. Male: Gonocoxites 9 as huge plates, apically rounded, gonostyli 9 not discernible, gonapophyses 9 processus-like; ectoproct broadly rounded. Complex of gonocoxite + gonostylus + gonapophysis 10 partly amalgamated with sternite 9, forming i) a pseudoapex of the latter which is deeply forked and ii) a paired hook. Gonocoxites 11 fused into a broad plate with a large median tooth (fused gonostyli 11?).

Female

Forewing length 7.8–9.0 mm, hindwing length 6.8–8.0 mm.

Fused gonocoxites 8 forming a broad trapezoid sclerite; gonapophyses 8 fused to triangular sclerite; gonocoxites 9 club-shaped, without distinct gonostylus.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): “Australia, Vic. Aucheron R. Feb.1987 Zwick” (CSIRO).

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).

Distribution

Australia (Victoria, NSW).

Nipponeurorthus Nakahara, 1958

Nipponeurorthus Nakahara, 1958: 25 (odescr) [Type species: Nipponeurorthus pallidinervis Nakahara, 1958: 25, by original designation].

Nipponeurorthus Nakahara: Hayashi 2005 (list, distr, figs); U. Aspöck and H. Aspöck 2007 (fig: distrmap); U. Aspöck and H. Aspöck 2010a (fig: distrmap); Liu et al. 2014 (overview, fig: distrmap).

Diagnosis

Adults of small body-size; male forewing length 6–10 mm. Body coloration generally yellow. Forewings 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.

Nipponeurorthus damingshanicus Liu, H. Aspöck & U. Aspöck, 2014

Figs 4c; 8a–f; 16

Nipponeurorthus damingshanicus Liu, H. Aspöck & U. Aspöck, 2014: 225 (odescr, key, figs: wings, gs male, female, distrmap).

Type locality

China (Guangxi: Mt. Damingshan).

Male

Body length 4.5 mm; forewing length 7.7 mm, hindwing length 7.1 mm.

Head pale yellow. Antennae pale yellow. Mouthparts yellow; mandibles with brownish tips.

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.

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.

Female

Body length 5.3–5.6 mm; forewing length 8.1–8.2 mm, hindwing length 7.1–7.3 mm.

Fused gonocoxites 8 about twice as long as tergite 8, flatly and roundly plate-like, with posterior portion feebly sclerotized. Gonapophyses 8 subtriangular, largely covered by gonocoxite 8, lateral margins distinctly sclerotized. Bursa copulatrix comprising a large and arcuate sclerotized sclerite, which is nearly as long as gonocoxite 8.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): “CHINA, Guangxi, Wuming, Mt. Damingshan [23°29’N, 108°26’E, 1257 m], 25.V.2011, Tingting Zhang” (CAU).

Biology and ecology

Adults have been taken in May. The known vertical distribution is 1257 m. The larva is unknown.

Distribution

China (Guangxi).

Figure 8. 

Genital segments of Nipponeurorthus damingshanicus Liu, H. Aspöck & U. Aspöck. a–d Male holotype: a lateral b caudal c dorsal d ventral e–f Female paratype: e genital segments, lateral f ventral. Scale bar: 0.5 mm.

Nipponeurorthus fasciatus Nakahara, 1958

Figs 4a; 9a–f; 16

Nipponeurorthus fasciatus Nakahara, 1958: 28 (odescr, figs: wings, gs male): Nakahara 1966 (distr); Hayashi 2005 (list, distr, figs); U. Aspöck and H. Aspöck 2007 (fig: distrmap); U. Aspöck and H. Aspöck 2008b (distrmap); Liu et al. 2014 (key, fig: distrmap).

Type locality

China (Taiwan: Urai).

Male

Forewing length 7.6–7.7 mm, hindwing length 6.7–7.2 mm.

Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.

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.

Abdomen yellow, dorsally largely tinged with pale reddish brown. 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 with a feebly produced subdistal projection. Ectoproct broad, directed posteroventrad, and slightly concaved medially on posterior margin. Complex of gonocoxites + gonostyli + gonapophyses 10 rather small; lateral arms much longer than distal projections, which are slenderly digitiform and parallelly directed dorsad. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as posteriorly bifurcated sclerite.

Female

Forewing length 11.7 mm, hindwing length 10.8 mm.

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.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): China, “Urai Formosa, June 20, 1922 (Dr. Kichizo Takeuchi)” (NSMT).

Biology and ecology

Adults have been taken from April–June. The known vertical distribution is 1100 m.

Distribution

China (Taiwan).

Figure 9. 

Genital segments of Nipponeurorthus spp. a–f Nipponeurorthus fasciatus Nakahara, a–c male: a lateral b ventral c dorsal d–f female: d lateral e gonocoxites 8 and gonapophyses 8, ventral f bursa copulatrix g–i Nipponeurorthus flinti U. Aspöck & H. Aspöck, male holotype: g ventral h lateral i dorsal. Scale bars: 0.5 mm.

Nipponeurorthus flinti U. Aspöck & H. Aspöck, 2008

Figs 4e; 9g–i; 16

Nipponeurorthus flinti U. Aspöck & H. Aspöck, 2008b: 818 (odescr, figs: wings, gs male, distrmap); Liu et al. 2014 (key, fig: distrmap).

In the heading of the original description (U. Aspöck and H. Aspöck 2008b: page 818) it is erroneously written “Austroneurorthus flinti”. This is a lapsus calami.

Type locality

Japan (Okinawa: Yonagawa, Yona).

Male

Body length 5.0–5.3 mm; forewing length 6.5–8.5 mm, hindwing length 6.0–6.6 mm.

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.

Female

Unknown.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): Japan, “Okinawa: Kunigami-gun upper Yonagawa, Yona 26°45.0’N, 12”8°13.3'E, 25 March 1997 O.S. Flint, Jr.” (NMNH).

Biology and ecology

Adults have been taken in March and May. No data concerning vertical distribution are available.

Distribution

Japan (Okinawa, Amamioshima).

Nipponeurorthus furcatus Liu, H. Aspöck & U. Aspöck, 2014

Figs 4d; 10a–c; 16

Nipponeurorthus furcatus Liu, H. Aspöck & U. Aspöck, 2014: 229 (odescr, key, figs: wings, gs male, distrmap).

Type locality

China (Yunnan: Lvchun).

Male

Body length 4.0 mm; forewing length 7.1–7.4 mm, hindwing length 6.5–6.9 mm.

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 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.

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.

Female

Unknown.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): “CHINA, Yunnan, Lvchun, Qimaba, Dapingzhang [22°50’N, 102°13’E], 1600 m, 21.VII.2013, Yang Zhao” (CAU).

Biology and ecology

Adults have been taken in July. The known vertical distribution is 1600 m. The larva is unknown.

Distribution

China (Yunnan).

Figure 10. 

Genital segments of Nipponeurorthus spp. a–c. Nipponeurorthus furcatus Liu, H. Aspöck & U. Aspöck, male holotype, a: lateral; b: dorsal; c: ventral; d–h. Nipponeurorthus fuscinervis (Nakahara), d-e: male, d: lateral, e: ventral, f–h: female, f: lateral; g: bursa copulatrix; h: gonocoxites 8 and gonapophyses 8, ventral. Scale bars: 0.5 mm.

Nipponeurorthus fuscinervis (Nakahara, 1915)

Figs 2c; 10d–h; 16

Neurorthus fuscinervis Nakahara, 1915: 16 (odescr, figs: gs female).

Nipponeurorthus fuscinervis: Nakahara 1958 (charact, figs: wing, gs male, female); Hayashi 2005 (list, distr, figs); U. Aspöck and H. Aspöck 2008b (fig: distrmap); Liu et al. 2014 (key, fig: distrmap).

Type locality

Japan (Kyoto: Mt. Atago).

Male

Forewing length 8.9–9.3 mm, hindwing length 7.5–7.8 mm.

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.

Female

Forewing length 8.8 mm, hindwing length 7.6 mm.

Fused gonocoxites 8 about 2.0 times as long as tergite 8, flatly plate-like. Gonapophyses 8 subtrapezoidal, largely covered by gonocoxite 8, lateral margins distinctly sclerotized. Bursa copulatrix sac-like, nearly hexagonal in ventral view, slightly longer than tergite 8; distal portion internally with an ovoid sclerotized area, terminally curved dorsad in lateral view.

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).

Nipponeurorthus multilineatus Nakahara, 1966

Figs 4h; 11a–f; 16

Nipponeurorthus multilineatus Nakahara, 1966: 204 (odescr, figs: wing, gs male, female); U. Aspöck and H. Aspöck 2008b (fig: distrmap); Liu et al. 2014 (key, fig: distrmap).

Type locality

China (Taiwan: Ilan).

Male

Forewing length 8.3–8.9 mm, hindwing length 7.2–7.6 mm.

Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.

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.

Abdomen yellow, dorsally purplish brown. Gonocoxite 9 robust on proximal half, with a small hairy tubercle on inner surface; distal half strongly incurved, ventrally with a subtriangular lobe; gonostylus 9 spinous and unforked. Ectoproct broad, directed posteroventrad, with posterior margin slightly concaved. Complex of gonocoxites + gonostyli + gonapophyses 10 present as a pair of slender straight lobes, which are directed posterodorsally. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as posteriorly bifurcated sclerite.

Female

Forewing length 9.7-9.9 mm, hindwing length 8.3-8.8 mm.

Fused gonocoxites 8 about 1.5 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, subquadrate in ventral view, nearly as long as tergite 8; distal portion internally with an ovoid sclerotized area, terminally curved dorsad in lateral view.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): China, “Ilan, Taipei Hsien, Formosa, April 16 1965 (Hirashima)” (NSMT).

Biology and ecology

Adults have been taken in April. No data concerning the vertical distribution are available. The larva is unknown.

Distribution

China (Taiwan).

Figure 11. 

Genital segments of Nipponeurorthus spp. a–f Nipponeurorthus multilineatus Nakahara a–c male: a lateral b ventral c dorsal d–f female: d lateral e gonocoxites 8 and gonapophyses 8, ventral f bursa copulatrix, ventral g–j Nipponeurorthus pallidinervis Nakahara g–i male paratype: g lateral h ventral i dorsocaudal j–l female paratype: j lateral k gonocoxites 8 and gonapophyses 8, ventral l bursa copulatrix. Abbreviations. b – bursa copulatrix; e – ectoproct; g – ring of glands; gp – gonapophysis; gs – gonostylus; gx – gonocoxite; S – sternite; T – tergite. Scale bars: 0.5 mm.

Nipponeurorthus pallidinervis Nakahara, 1958

Figs 4f–g; 11g–k; 16

Nipponeurorthus pallidinervis Nakahara, 1958: 25 (odescr, figs: wing, gs male, female); Kuwayama 1962 (fig. body, wings); Zwick 1967 (figs: gs female); Hayashi 2005 (list, distr, figs); U. Aspöck and H. Aspöck 2008a (compmorphol, figs: gs male); U. Aspöck and H. Aspöck 2008b (fig: distrmap); Liu et al. 2014 (key, fig: distrmap).

Type locality

Japan (Hokkaido: Jozankei).

Male

Forewing length 8.8–9.8 mm, hindwing length 7.4–8.6 mm.

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.

Female

Forewing length 9.1–11.4 mm, hindwing length 8.0–9.9 mm.

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, slightly longer than tergite 8, with distal portion laterally expanded in ventral view, marginally and internally with several sclerotized bands.

Specimens examined and records published

Supplementary material 1. Holotype male (by original designation): Japan, “Jozankei, Hokkaido, July 17–18, 1956, Waro Nakahara” (NSMT).

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).

Nipponeurorthus punctatus (Nakahara, 1915)

(Figs 4i; 12a–e; 16)

Neurorthus punctatus Nakahara, 1915: 15 (odescr, figs wings): Navás 1935 (mon, fig: wing).

Nipponeurorthus punctatus (Nakahara, 1915): Okamoto and Kuwayama 1932 (fig. body, wings); Nakahara 1958 (figs: wings, gs male, female); Zwick 1967 (figs: gs female); Hayashi 2005 (list, distr, figs); U. Aspöck and H. Aspöck 2008b (fig: distrmap); Liu et al. 2014 (key, fig: distrmap).

Type locality

Japan (Honshu: Tottori, or Kyoto: Mt. Atago, or Osaka: Mt. Minomo) [A lectotype should be designated, however, the syntypes (from the above mentioned localities) are unavailable presently].

Male

Forewing length 7.1–7.4 mm, hindwing length 6.2–6.5 mm.

Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.

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.

Abdomen yellow, dorsally slightly darker. Gonocoxite 9 robust on proximal half; distal half strongly incurved, ventrally with a short digitiform projection, which bears several spines; gonostylus 9 spinous and forked into a triangular subdistal projection. Ectoproct broad, directed posteroventrad. Complex of gonocoxites + gonostyli + gonapophyses 10 with sinuate lateral arms, which are inflated posterolaterally; distal projections slenderly digitiformed, straightly directed posteriad. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as posteriorly bifurcated sclerite.

Female

Forewing length 7.7–8.9 mm, hindwing length 7.2–7.9 mm.

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.

Distribution

Japan (Honshu, Hokkaido, Kyushu).

Figure 12. 

Nipponeurorthus spp. a–e Nipponeurorthus punctatus (Nakahara) a–b male genital segments: a lateral b ventral c–e female genital segments: c lateral d gonocoxites 8 and gonapophyses 8, ventral e bursa copulatrix f Nipponeurorthus qinicus Yang in Chen, male holotype, habitus drawing (adapted from Yang in Chen 1998) g–i Nipponeurorthus tianmushanus Yang & Gao, male: g wings h genital segments, lateral i caudal (adapted from Yang and Gao 2001) j–m Nipponeurorthus tinctipennis Nakahara, male: j forewing k–m genital sclerites (adapted from Nakahara 1958). Scale bar: 0.5 mm (a–e).

Nipponeurorthus qinicus Yang in Chen, 1998

Figs 12f; 16

Nipponeurorthus qinicus Yang in Chen, 1998: 105 (odescr, figs: habitus); Liu et al. 2014 (key, fig: distrmap).

Type locality

China (Shaanxi: Ankang).

Male

Body length 7.0 mm; forewing length 9.5 mm, hindwing length 8.0 mm.

Head yellow. Antennae yellow but gradually darkened toward apex.

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.

Abdomen yellow. Gonocoxite 9 strongly curved distad. Ectoproct broad, slightly concaved on posterior margin. Complex of gonocoxites + gonostyli + gonapophyses 10 present as a pair of hook-like lobes. Gonostyli 11 present as posteriorly bifurcated sclerite.

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.

Distribution

China (Shaanxi).

Nipponeurorthus tianmushanus Yang & Gao, 2001

Figs 12g–i; 16

Nipponeurorthus tianmushanus Yang & Gao, 2001: 308 (odescr, figs: wings, gs male): Liu et al. 2014 (key, fig: distrmap).

Type locality

China (Zhejiang: Tianmushan).

Male

Body length 7.0 mm; forewing length 8.0 mm, hindwing length 7.0 mm.

Head yellow. Antennae yellowish brown, with several terminal flagellomeres dark brown.

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).

Nipponeurorthus tinctipennis Nakahara, 1958

Figs 4j; 12j–m; 16

Nipponeurorthus tinctipennis Nakahara, 1958: 27 (odescr, figs: wing, gs male, female); Hayashi 2005 (list, distr, figs); U. Aspöck and H. Aspöck 2008b (fig: distrmap); Liu et al. 2014 (key, fig: distrmap).

Type locality

Japan (Yakushima Island: Hananoegou and Muromidake).

Male

Forewing length 9.1 mm, hindwing length 8.0 mm.

Head yellow. Antennae yellow. Mouthparts yellow; mandibles with brownish tips.

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.

Abdomen yellow, dorsally purplish brown. Gonocoxite 9 robust on proximal half, distal half strongly incurved; gonostylus 9 spinous. Complex of gonocoxites + gonostyli + gonapophyses 10 present as a pair of slender lobes, which are inflated distad and bear a tooth-like processus. Gonocoxites 11 present as a simple, transverse, sclerotized band; gonostyli 11 present as posteriorly bifurcated sclerite.

Female

Forewing length 10.0 mm, hindwing length 9.0 mm.

Fused gonocoxites 8 flatly plate-like. Gonapophyses 8 subtriangular, largely covered by gonocoxite 8, lateral margins distinctly sclerotized.

Specimens examined and records published

Supplementary material 1. Holotype male (by original description), Japan, “Hananoegou, Yakushima, 12 July 1954, Yosihiko Kurosawa” (NSMT).

Biology and ecology

The adult has been taken in July. The known vertical distribution is 1800 m. The larva is unknown.

Distribution

Japan (Yakushima Island).

Sinoneurorthus Liu, H. Aspöck & U. Aspöck, 2012

Sinoneurorthus Liu, H. Aspöck & U. Aspöck, 2012: 132 (odescr) [Type species: Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, 2012: 133, by original designation].

Diagnosis

Adults of medium body size; female forewing length 12-13 mm. Body coloration reddish orange. Wings slightly leathery, smoky brown. Longitudinal veins with dense branches, leaving small bifurcated or trifurcated forks marginally. Female fused gonocoxite 8 flatly and roundly plate-like; gonocoxites 9 narrowly foliate, with ovoid gonostyli; bursa copulatrix distinctly shaped and sclerotized.

Distribution

China.

Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, 2012

Figs 2d; 4k; 13a–e; 16

Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, 2012: 133 (odescr, figs: adult, wings, gs female, distrmap).

Type locality

China (Yunnan: Xiaocaoba).

Female

Body length 6.9 mm; forewing length 12.6 mm, hindwing length 11.0 mm.

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.

Abdomen reddish orange. Fused gonocoxites 8 about twice as long as tergite 8, flatly and roundly plate-like. Gonapophyses 8 subtrapezoidal, proximal half covered by gonocoxites 8, lateral margins distinctly sclerotized. Bursa copulatrix comprising an ovoid sclerotized sclerite, with a pair of cone-shaped hollow processes directed ventrad.

Male

Unknown.

Specimens examined and records published

Supplementary material 1. Holotype female (by original designation): “CHINA: Yunnan Province, Zhaotong City, Yiliang County, Xiaocaoba, 27°50.079N, 104°17.554E, 1715 m, 2009.V.28, Liangming Cao leg.” (CAU).

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.

Distribution

China (Yunnan).

Figure 13. 

Sinoneurorthus yunnanicus Liu, H. Aspöck & U. Aspöck, female holotype a wings b genital segments, lateral c same, dorsal d same, ventral e gonocoxites 9, lateral f bursa copulatrix, lateral. Abbreviations. A – Analis; –: Costa; CuA – Cubitus anterior; CuP – Cubitus posterior; J – Jugal vein; MA – Media anterior; MP – Media posterior; R – Radius; Rs – Radial sector; Sc – Subcosta. b – bursa copulatrix; e – ectoproct; gp – gonapophysis; gs – gonostylus; gx – gonocoxite; S – sternite; T – tergite. Scale bar: 1.0 mm (a) and 0.5 mm (b–f).

Figure 14. 

Distribution map of the species of Nevrorthus.

Figure 15. 

Distribution map of the species of Austroneurorthus.

Figure 16. 

Distribution map of the species of Nipponeurorthus and Sinoneurorthus.

Figure 17. 

Distribution map of the family Nevrorthidae.

Key to extant genera of Nevrorthidae

1 Wing membrane slightly leathery (only female known) (Fig. 2d) Sinoneurorthus
Wing membrane soft 2
2 Males: Segment 7 enlarged (Figs 6d, e), distribution restricted to Mediterranean (and submediterranean) regions Nevrorthus
Males: Segment 7 not enlarged 3
3 Males: Complex of gonocoxites, gonostyli, gonapophyses 11 forming a transverse sclerite (Fig. 7e), distribution restricted to Australia Austroneurorthus
Males: Complex of gonocoxites, gonostyli, gonapophyses 11 with a small median fork (Fig. 11i), distribution restricted to Eastern Asia Nipponeurorthus

Key to extinct genera of Nevrorthidae (all from the Eocene Baltic amber) (see Wichard 2016).

Key to species of Nevrorthus (males)

4 Forewing without shadows on cross veins (Figs 3a, d) 2
Forewing with shadows on cross veins (Figs 3b, e) 3
5 Scapus and pedicellus yellowish, pseudoapex of sternite 9 deeply forked (Fig. 6b) N. iridipennis
Scapus and pedicellus dark brown, pseudoapex of sternite 9 unforked (Fig. 6c N. apatelios
6 Flagellum of antennae uniformly yellowish brownish, pseudoapex of sternite 9 deeply grooved (Figs 6a, e) 4
Flagellum of antennae slightly darker in distal third, pseudoapex of sternite 9 distally sinuate (Fig. 3f) N. reconditus
7 Gonocoxites 11 forming a triangle (Fig. 6a) N. fallax
– Gonocoxites 11 forming a bar (Fig. 6e) N. hannibal

Key to species of Austroneurorthus (males)

1 Forewing with intensive shadows around crossveins (Fig. 3h), femora of all three legs in males with dark orange coloured oval sclerite; pseudoapex of sternite 9 rounded (Fig. 7f) A. horstaspoecki
Forewing without shadows around crossveins (Fig. 3g), femora of males without dark orange sclerites; pseudoapex of sternite 9 deeply forked (Fig. 7b) A. brunneipennis

Key to species of Nipponeurorthus

See Liu et al. (2014)

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.

Figure 18. 

Strict consensus tree of 40 MPTs generated from the refined data matrix. Bootstrap support values are shown at nodes. Only unambiguous character changes are shown. Black circles represent unique changes, white circles represent homoplasious changes. The symbol “†” indicates extinct genus.

Discussion

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 so-called “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) has been discussed for Mediterranean species (H. Aspöck et al. 2001, U. Aspöck and H. Aspöck 2010a), all of them constituting the genus Nevrorthus. Refugial centres that would be relevant to Nipponeurorthus and Sinoneurorthus are poorly understood (Liu et al. 2012, 2014). The biogeographic origin of Austroneurorthus remains enigmatic (U. Aspöck 2004).

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 so-called 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.

Acknowledgements

We want to express our cordial thanks to David Britton (Sydney), Josefine Cardale (Canberra), Oliver Flint and David Furth (Washington DC), Niels P. Kristensen† (Copenhagen), Wolfram Mey (Berlin), John Oswald (College 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) and the Beijing Natural Science Foundation (No. 5162016). We also acknowledge the Museum für Naturkunde, Berlin, for the support during the publication of this article.

References

  • Aspöck H, Aspöck U, Hölzel H (1977) Neurorthus apatelios n. sp. – eine verkannte europäische Neurorthiden-Species (Neuroptera: Planipennia). Entomologische Zeitschrift, Frankfurt am Main 87: 53–57.
  • Aspöck H, Aspöck U, Hölzel H (1978) Megaloptera et Planipennia. In: Illies J (Ed.) Limnofauna Europaea. 2nd edition. Gustav Fischer Verlag, Stuttgart, 329–332.
  • Aspöck H, Aspöck U, Hölzel H [unter Mitarbeit von H. Rausch] (1980) Die Neuropteren Europas. 2 vols. Goecke & Evers, Krefeld. 495 pp., 355 pp.
  • Aspöck H, Hölzel H (1996) The Neuropteroidea of North Africa, Mediterranean Asia and of Europe: a comparative review (Insecta). In Canard M, Aspöck H, Mansell MW (Eds) Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neueropterology (2–6 May 1994, Cairo, Egypt). Toulouse, France, 31–86.
  • Aspöck H, Hölzel H, Aspöck U (2001) Kommentierter Katalog der Neuropterida (Insecta: Raphidioptera, Megaloptera, Neuroptera) der Westpaläarktis. Denisia 2, 1–606.
  • Aspöck U (1992) Crucial points in the phylogeny of the Neuroptera (Insecta). In Canard M, Aspöck H, Mansell MW (eds). Current Research in Neuropterology. Proceedings of the Fourth International Symposium on Neuropterology (24–27 June 1991, Bagnères-de-Luchon, Haute-Garonne, France). Toulouse, 63–73.
  • Aspöck U (1993) Geklärtes und Ungeklärtes im System der Neuroptera (Insecta: Holometabola). Mitteilungen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie 8: 451–456.
  • Aspöck U (1995) Neue Hypothesen zum System der Neuropterida. Mitteilungen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie 10: 633–636.
  • Aspöck U (2004) Austroneurorthus horstaspoecki nov. spec. – eine neue Art der Familie Nevrorthidae aus Australien (Neuropterida: Neuroptera). Denisia 13: 177–182.
  • Aspöck U, Aspöck H (1983) Über das Vorkommen von Neurorthus Costa in Nordafrika (Neuropteroidea, Planipennia, Neurorthidae). Nachrichtenblatt der Bayerischen Entomologen 32: 48–51.
  • Aspöck U, Aspöck H (1994) Paradoxe Verbreitungsbilder bei Neuropteroidea (Insecta: Raphidioptera, Neuroptera). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 46: 30–44.
  • Aspöck U, Aspöck H (1999) Kamelhälse, Schlammfliegen, Ameisenlöwen ... Wer sind sie? (Insecta: Neuropterida: Raphidioptera, Megaloptera, Neuroptera). Stapfia 60: 1–34.
  • Aspöck U, Aspöck H (2007) Verbliebene Vielfalt vergangener Blüte. Zur Evolution, Phylogenie und Biodiversität der Neuropterida (Insecta: Endopterygota). Denisia, 20: 451–516.
  • Aspöck U, Aspöck H (2008b) Nipponeurorthus flinti nov. sp. – eine neue Art der Familie Nevrorthidae von der Insel Okinawa (Neuropterida: Neuroptera). Linzer biologische Beiträge 40: 817–825.
  • Aspöck U, Aspöck H (2010a) Erobern Nevrorthidae Mitteleuropa? Eine biogeographische Provokation (Neuroptera: Neuropterida). Denisia 29: 25–35.
  • Aspöck U, Aspöck H (2010b) Landmarks towards a phylogeny-based classification of the Neuropterida (Insecta: Endopterygota). In: Devetak D, Lipovšek S, Arnett AE (Eds) Proceedings of the Tenth International Symposium on Neuropterology (22–25 June 2008, Piran, Slovenia). University of Maribor, Maribor, 67–74.
  • Aspöck U, Plant JD, Nemeschkal HL (2001) Cladistic analysis of Neuroptera and their systematic position within the Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology 26: 73–86. https://doi.org/10.1046/j.1365-3113.2001.00136.x
  • Berendt GC (1845–1856) Die im Bernstein befindlichen organischen reste der Vorwelt, gesammelt in Verbindung mit mehreren bearbeitet und herausgegeben von Dr. Georg Carl Berendt. 2 Bd. Berlin. 374 pp.
  • Beutel RG, Friedrich F (2008) Comparative study of larval head structures of Megaloptera (Hexapoda). European Journal of Entomology 105: 917–938. https://doi.org/10.14411/eje.2008.119
  • Chen SC (1998) Pictorial handbook of rare and precious insects in China. China Forestry Publishing House, Beijing, 332 pp.
  • Costa A (1863) Nuovi studii sulla entomologia della Calabria ulteriore. Atti della Reale Accademia delle Scienze Fisiche e Matematiche di Napoli 1: 80 pp. https://doi.org/10.5962/bhl.title.34236
  • Devetak D (1992) Present knowledge of the Megaloptera, Raphidioptera and Neuroptera of Yugoslavia (Insecta: Neuropteroidea). In Canard M, Aspöck H, Mansell MW (Eds) Current Research in Neuropterology, Proceedings of the Fourth International Symposium on Neuropterology (24–27 June 1991, Bagnères de-Luchon, Haute-Garonne, France). Toulouse, 107–118.
  • Devetak D, Jakšić PN (2003) Neuroptera of Kosovo and Metohija (Serbia). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 55: 45–53.
  • Devetak D, Klokočovnik V (2016) The feeding biology of adult lacewings (Neuroptera): a review. Trends in Entomology 12: 30–42.
  • Esben-Petersen P (1913) Addition to the knowledge of the neuropterous insect fauna of Corsica II. Entomologiske Meddelelser 10: 20–28.
  • Esben-Petersen P (1929) Australian Neuroptera. VI. The Queensland Naturalist VII: 31–35. Gaumont J (1968) Nouvelles observations sur la disparition du cryptonéphridisme chez les larves aquatiques de Planipennes. Comptes rendus hebdomadaires des séances de la Académie des Sciences Paris 266: 2097–2099.
  • Gaumont J (1976) La appareil digestiv des larves de Planipennes. Annales des Sciences Naturelles, Zoologie et Biologie Animale (12)18: 145–250.
  • Gavira O, Sánchez S, Carrasco P, Ripoll J, Solís S (2012) Presence of the family Nevrorthidae (Insecta: Neuroptera) in the Iberian Peninsula. Boletin de la Sociedad Entomológica Aragonesa 51: 217–220.
  • Gepp J (1984) Erforschungsstand der Neuropteren-Larven der Erde (mit einem Schlüssel zur Larvaldiagnose der Familien, einer Übersicht von 340 beschriebenen Larven und 600 Literaturzitaten). In: Gepp J, Aspöck H, Hölzel H (Eds) Progress in World`s Neuropterology. Proceedings of the First International Symposium on Neuropterology (September 1980, Graz, Austria). Graz, 183–239.
  • Güsten R (1996) A review of epidermal glands in the order Neuroptera (Insecta). In: Canard M, Aspöck H, Mansell MW (Eds) Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology (2–6 May 1994, Cairo, Egypt). Toulouse, 129–146.
  • Hagen HA (1864) Névroptères (non Odonates) de la Corse, recueillis par M. E. Bellier de la Chavignerie en 1860 et 1861. Annales de la Société Entomologique de France (4)4: 38–45.
  • Handlirsch A (1906–1908) Die fossilen Insekten und die Phylogenie der rezenten Formen – ein Handbuch für Paläontologen und Zoologen. 2 Bd. Engelmann, Leipzig, 1430 pp.
  • Hayashi F (2005) Neuroptera. In Kawai T, Tanida K (eds). Aquatic insects of Japan: Manual with keys and illustrations. Tokai University Press, Kanagawa, 387–392. [in Japanese]
  • Henry CS (1982) Neuroptera. In: McGraw H (Ed.) Synopsis and Classification of Living Orders. New York, 470–482.
  • Iori A, Kathirithamby J, Letardi A, Pantaleoni RA, Principi MM (1995) Neuropteroidea (Megaloptera, Raphidioptera, Planipennia), Mecoptera, Siphonaptera, Strepsiptera. In: Minelli A, Ruffo S, La Posta S (Eds) Checklist delle Specie della Fauna Italiana 62, 1–20.
  • Jones R, Devetak D (2009) First record of Nevrorthidae from Slovenia. Acta Entomologica Slovenica 17: 99–106.
  • Kimmins DE (1930) A list of the Corsican Ephemeroptera and Neuroptera. Eos: Revista Española de Entomología 6: 185–190.
  • Klapálek F (1917) Über die von Herrn Prof. A Hetschko in Korsika gesammelten Neuropteroiden nebst Bemerkungen über einige ungenügend bekannte Arten. Wiener Entomologische Zeitung 36: 193–208. https://doi.org/10.5962/bhl.part.12934
  • Krüger L (1923) Neuroptera succinica baltica. Die im baltischen Bernstein eingeschlossenen Neuroptera des Westpreussischen Provinzial-Museums (heute Museum für Naturkunde und Vorgeschichte) in Danzig. Stettiner Entomologische Zeitung 84: 68–92.
  • Kuwayama S (1962) A revisional synopsis of the Neuroptera in Japan. Pacific Insects 4: 325–412.
  • Lattin G de (1967) Grundriss der Zoogeographie. G. Fischer Verlag, Stuttgart, 602 pp.
  • Leraut P (1981) Liste des Planipennes de France (Neuroptera). Bulletin de la Société Entomologique de France 85: 237–253.
  • Lestage JA (1924) Où faut-il placer les Neurorthus? Annales de la Société Entomologique de Belgique 64: 65–72.
  • Letardi A (1994) Dati sulla distribuzione italiana dei Neurotteri s.l. (Neuropteroidea). Atti XVII Congresso Nazionale Italiano di Entomologia (Udine 13–18 June 1994), 203–206.
  • Letardi A, Aspöck U, Aspöck H, Pantaleoni RA (2006) Nevrorthus apatelios Aspöck H et Aspöck U et Hölzel H, 1977 (Neuroptera Nevrorthidae) nelle Prealpi Friulane. Revista del Museo Civico di Scienze Naturali “E. Caffi” 24[2005]: 91–92.
  • Letardi A, Pantaleoni RA (1996) I neurotteroidei W-Paleartici della collezione del Museo di Zoologia dell’Università di Roma (Neuropteroidea). Fragmenta entomologica, Roma 28: 277–305.
  • Letardi A, Thierry D, Tillier P, Canard M (2008) Mise á jour de la faune des Neuropterida de Corse (Raphidioptera & Neuroptera). Revue de l’Association Roussillonnaise d’Entomologie 17: 95–105.
  • Liu XY, Aspöck H, Aspöck U (2012) Sinoneurorthus yunnanicus n. gen. et n. sp. – a spectacular new species and genus of Nevrorthidae (Insecta: Neuroptera) from China, with phylogenetic and biogeographical implications. Aquatic Insects 34: 131–141. https://doi.org/10.1080/01650424.2012.718086
  • Makarkin VN (2016) The neuropteran assemblage (Insecta) of the mid-Cretaceous Burmese amber confirms transitional character of its biota. In: Dzyuba OS, Pestchevitskaya EB, Shurygin BN (Eds) Cretaceous Ecosystems and Their Responses to Paleoenvironmental Changes in Asia and the Western Pacific: Short papers for the Fourth International Symposium of IGCP Project 608, Novosibirsk, August 15-20, 2016. IPGG SB RAS, Novosibirsk, 27–29.
  • Makarkin VN, Perkovsky EE (2009) Rophalis relicta Hagen (Neuroptera, Nevrorthidae) in Late Eocene Rovno amber, with a discussion of the taxonomic status of the genus. Denisia 26: 137–144.
  • Malicky H (1984) Ein Beitrag zur Autökologie und Bionomie der aquatischen Netzflüglergattung Neurorthus (Insecta, Neuroptera, Neurorthidae). Archiv für Hydrobiologie (und Planktonkunde) Stuttgart 101: 231–246.
  • McLachlan R (1881) Sartena (Hagen, 1864) = Neurorthus (Costa, 1863). Entomologist’s Monthly Magazine 18: 89.
  • McLachlan R (1898) Neuroptera-Planipennia collected in Algeria by the Rev. A. E. Eaton. Transactions of the [Royal] Entomological Society of London 46: 151–168.
  • Monserrat VJ (1977) A systematic and alphabetic list of Neurorthidae and Sisyridae (Neuroptera). Nouvelle Revue d’Entomologie 7: 91–96.
  • Monserrat VJ (1985) Lista de los tipos de Mecoptera y Neuroptera (Insecta) de la colección Navás L, depositados en el Museo de Zoología de Barcelona. Miscellánia Zoológica 9: 233–243.
  • Monserrat VJ (2005) Nuevos datos sobre algunas peqeñas familias de neurópteros (Insecta: Neuroptera: Nevrorthidae, Osmylidae, Sisyridae, Dilaridae). Heteropterus Revista de Entomología 5: 1–26.
  • Monserrat VJ, Triviño V (2013) Atlas de los neurópteros de la Península Ibérica e Islas Baleares (Insecta, Neuroptera: Megaloptera, Raphidioptera, Planipennia). Monografias de la Sociedad Entomológica Aragonesa 13: 1–154.
  • Nakahara W (1915) On the Hemerobiinae of Japan. Annotationes Zoologicae Japonenses 9: 11–48.
  • Nakahara W (1958) The Neurorthinae, a new subfamily of the Sisyridae (Neuroptera). Mushi 32: 19–32.
  • Nakahara W (1966) Hemerobiidae, Sisyridae and Osmylidae of Formosa and Ryukyu Islands (Neuroptera). Kontyû 34: 193–207.
  • Navás L (1935) Monografía de la familia de los Sisíridos (Insectos Neurópteros). Memorias de la Real Academia de Ciencias Exactas, Fisico-Quimicas y Naturales de Zaragoza 4: 1–87.
  • Nel A, Jarzembowski E (1997) New fossil Sisyridae and Nevrorthidae (Insecta: Neuroptera) from Eocene Baltic amber and Upper Miocene of France. European Journal of Entomology 94: 287–294.
  • New TR (1978) Note on the habitat of the presumed larva of Austroneurorthus (Neuroptera, Neurorthidae). Australian Entomological Magazine 5: 9.
  • New TR (1986) A review of the biology of Neuroptera Planipennia. Neuroptera International 1(Suppl. Series): 1–57.
  • New TR (1989) Planipennia, Lacewings. Handbuch der Zoologie. Vol. 4 (Arthropoda: Insecta), Part 30. Walter de Gruyter, Berlin: 132 pp.
  • New TR (1991) Neuroptera (Lacewings). In: Naumann ID (Ed.) The Insects of Australia. 2nd Edition. Vol. 1 Melbourne University Press, Melbourne, 525–542.
  • New TR (1996) Neuroptera. In: Wells A (Ed.) Zoological Catalogue of Australia. Vol. 28: Neuroptera, Strepsiptera, Mecoptera, Siphonaptera. Melbourne. CSIRO Publishing, Melbourne. 1–104, 184 (Appendix III), 199–216.
  • Nicoli Aldini R, Letardi A, Pantaleoni RA (2012) State of the art on Neuropterida of Sicily and Malta. Biodiversity Journal 3: 445–458.
  • Nixon KC (2002) WinClada ver. 1.00.08. Published by the author, Ithaca, NY.
  • Okamoto H, Kuwayama S (1932) Neuroptera. In: Esaki et al. (Eds) Iconographia insectorum Japonicorum (1st edn). Hokuryukan Co., Tokyo, 1524–1555.
  • Oswald JD, Penny ND (1991) Genus-group names of the Neuroptera, Megaloptera and Raphidioptera of the world. Occasional papers of the California Academy of Sciences 147. California Academy of Sciences, San Francisco, California, 94 pp.
  • Pantaleoni RA (1999) Neuropterida described by A. Costa with type designation. Deutsche Entomologische Zeitschrift 46: 249–261.
  • Pantaleoni RA (2005) Interpretation of Achille Costa’s data on Neuropterida. Bulletin of Insectology 58: 71–92.
  • Parfin SI, Gurney AB (1956) The Spongillaflies, with special reference to those of the western hemisphere (Sisyridae, Neuroptera). Proceedings of the United States National Museum 105: 421–529. https://doi.org/10.5479/si.00963801.105-3360.421
  • Pongrácz S (1923) Recésszárnyúak. Neuropteroiden. In Csiki E (Ed.) Állattani Kutatásai Albániában (Explorationes zoologicae ab E. Csiki in Albania peractae). XI. A. Magyar Tudományos Akadémia, Balkán-Kutatásainak Tudományos Erdményei. Vol. 1. Budapest, 143–166.
  • Popov A (1990) Beitrag zur Kenntnis der Neuropteren des Witoscha Gebirges. Fauna of Southwestern Bulgaria 3: 78–87. [In Bulgarian, summary in German]
  • Popov A (1991) Baum- und strauchbewohnende Neuropteren in Bulgarien. Acta zoologica bulgarica 41: 26–36.
  • Popov A (1992) Zoogeographical analysis of Neuropteroidea (Insecta) of the Balkan Peninsula. In: Canard M, Aspöck H, Mansell W (Eds) Current Research in Neuropterology. Proceedings of the Fourth International Symposium on Neuropterology (24-27 June 1991, Bagnères de-Luchon, Haute-Garonne, France). Toulouse, 319–330.
  • Popov A (1993) Raphidiopteren und Neuropteren aus Bulgarien in den Sammlungen des Nationalmuseums in Prag. Historia Naturalis Bulgarica 4: 16–28.
  • Popov A (2007) Distribution of the families of Neuroptera with low species diversity in Bulgaria. In: Pantaleoni RA, Letardi A, Corazza C (Eds) Proceedings of the Ninth International Symposium on Neuropterology (20–23 June 2005, Ferrara, Italy). Annali del Museo Civico di Storia Naturale di Ferrara 8[2005], 117–130.
  • Principi MM (1966) Contributi allo studio dei Neurotteri Italiani. XVIII. Neurotteri della Basilicata, della Calabria e della Sicilia. Memorie del Museo Civico di Storia Naturale di Verona 14: 363–388.
  • Rambur MP (1842) Histoire naturelle des insectes. Névroptères. Roret, Paris, 534 pp.
  • Randolf S, Zimmermann D, Aspöck U (2014) Head anatomy of adult Nevrorthus apatelios and basal splitting events in Neuroptera (Neuroptera: Nevrorthidae). Arthropod Systematics & Phylogeny 72: 111–136. https://doi.org/10.1016/j.asd.2013.07.004
  • Randolf S, Zimmermann D, Aspöck U (2013) Head anatomy of adult Sisyra terminalis (Insecta: Neuroptera: Sisyridae) – functional adaptations and phylogenetic implications. Arthropod Structure & Development 42: 565–582.
  • Riek EF (1970) Neuroptera. In The Insects of Australia. Melbourne University Press, Melbourne, 472–494.
  • Saure C (1989) Beitrag zur Kenntnis der Neuropterenfauna Jugoslawiens und Griechenlands (Insecta, Planipennia). Entomofauna 10: 33–43.
  • Starmühlner F (1982) Auf der Suche nach “lebenden Fossilien”. Zoologische Garten N. F. 52: 152–160.
  • Stelzl M (1992) Comparative studies on mouthparts and feeding habits of adult Raphidioptera and Neuroptera (Insecta: Neuropteroidea). In: Canard M, Aspöck H, Mansell W (Eds) Current Research in Neuropterology. Proceedings of the Fourth International Symposium on Neuropterology (24_27 June 1991, Bagnères de-Luchon, Haute-Garonne, France). Toulouse, 341–347.
  • Sziráki G (2008) A Nevrorthidae Nakahara, 1915 (Insecta: Neuroptera) Család elöfordulása Cserna-Hegységben. Acta Siculica 2008: 87–90.
  • Takahashi R (1942) Uncommon neuropterous larva. Zoological Magazine 54: 439–441.
  • Thenius E (2000) Lebende Fossilien: Oldtimer der Tier- und Pflanzenwelt – Zeugen der Vorzeit. Dr. Friedrich Pfeil, München, 228 pp.
  • Wachmann E, Saure C (1997) Netzflügler, Schlamm- und Kamelhalsfliegen: Beobachtung - Lebensweise. Naturbuch Verlag, Augsburg, 159 pp.
  • Wang YY, Liu XY, Garzón-Orduña IJ, Winterton SL, Yan Y, Aspöck U, Aspöck H, Yang D (2016) Mitochondrial phylogenomics illuminates the evolutionary history of Neuropterida. Cladistics 2016: https://doi.org/10.1111/cla.12186
  • Wedmann S, Makarkin VN, Weiterschan T, Hörnschemeyer T (2013) First fossil larvae of Berothidae (Neuroptera) from baltic amber, with notes on the biology and termitophily of the family. Zootaxa 3716: 236–258. https://doi.org/10.11646/zootaxa.3716.2.6
  • Wichard W (2014) Aquatische Neuropteren im Baltischen Bernstein. DGaaE-Nachrichten 28: 5.
  • Wichard W, Arens W, Eisenbeis G (1995) Atlas zur Biologie der Wasserinsekten. Gustav Fischer Verlag, Stuttgart, Jena & New York, 338 pp.
  • Wichard W, Gröhn C, Seredszus F (2009) Aquatic Insects in Baltic Amber. Verlag Kessel, Remagen, 336 pp.
  • Winterton SL, Hardy NB, Wiegmann BM (2010) On wings of lace: phylogeny and Bayesian divergence time estimates of Neuropterida (Insecta) based on morphological and molecular data. Systematic Entomology 35: 349–378.
  • Withycombe CL (1925) Some aspects of the biology and morphology of the Neuroptera. With special reference to the immature stages and their possible phylogenetic significance. Transactions of the Entomological Society of London 1924: 303–411. https://doi.org/10.1111/j.1365-2311.1925.tb03362.x
  • Xia FY, Yang GD, Zhang QQ, Shi GL, Wang B (2015) Amber: Lives through time and space. Science Press, Beijing, 197 pp.
  • Yang CK, Gao MY (2001) Neuroptera: Neurorthidae. In: Wu H, Pan CW (Eds) Insects of Tianmushan National Nature Reserve. Science Press, Beijing, 307–309.
  • Zelený J (1964) Ergebnisse der Albanien-Expedition 1961 des Deutschen Entomologischen Institutes. 24. Beitrag Neuroptera. Beiträge zur Entomologie 14: 323–336.
  • Zelený J (1971) Neuroptera, Megaloptera und Mecoptera aus Bulgarien. Acta Faunistica Entomologica Musei Nationalis Pragae 14: 153–164.
  • Zimmermann D, Randolf S, Metscher BD, Aspöck U (2011) The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta). Arthropod Structure & Development 40: 571–582. https://doi.org/10.1016/j.asd.2011.06.003
  • Zwick P (1967) Beschreibung der aquatischen Larve von Neurorthus fallax (Rambur) und Errichtung einer neuen Planipennierfamilie Neurorthidae fam. nov. Gewässer und Abwässer 44/45: 65–86.