Amphineurus Rhypholophus Amphineurus (A.) umbraticus Rhypholophus Systematics of Amphineurus (Rhamphoneurus Alexander) (Diptera: Tipuloidea: Limoniidae) Santos, Daubian Santos, Rodrigo dos Reis Ribeiro, Guilherme Cunha Arthropod Systematics & amp; Phylogeny 2022 2022-09-12 80 439 494 87DFG 9AA3C8A3-0567-51F0-A754-D36347DE9153 Skuse, 1890 Skuse 1890 Insecta Limoniidae Amphineurus CoL Animalia Amphineurus Diptera 0 439 Arthropoda genus   AmphineurusSkuse, 1890: 800 (as a subgenus of Rhypholophus). Type-species: Amphineurus (A.) umbraticusSkuse, 1890 (as Rhypholophus).  Diagnosis.   Amphineuruscan be diagnosed by the following combination of characters: gonocoxite bifid; wing with R2+3+4 aligned with R2+3. When there is an element R4+5 (and therefore, no R2+3), members of the genus are distinguished from other Chioneinaeby the closed discal cell.  Remarks. There is great diversity in the male terminalia of  Amphineurus(Figs 1, 2). There are many variations from the simplified pattern of A. (A.) patyaTheischinger (Fig. 1A) to the asymmetric disposition of A. (Nesormosia)(Fig. 2D). Therefore, defining a broad diagnosis for  Amphineurusis not so easy. Skuse (1890)defined  Amphineurusas a subgenus of  Rhypholophus. Skuse'sdefinition fits better to the current subgenus A. (Amphineurus)but does not portray the reality of other species of  Amphineurus. His diagnosis included the absence of the discal cell, the radial sector with four branches, "second longitudinal vein angulated", and hind femora one third longer than the intermediate pair. These features are non-exclusive or variable in the genus.   Figure 1.Schematic illustration of the male terminalia of representatives of  Amphineurus(dorsal view). A  A.( A.)  Amphineurus patya; B A. (A.) superbus; C A. (A.) pita; D A. (A.) umbraticus. - Abbreviations: ad spur, anterodorsal spur; aed, aedeagus; aed sh, sheath of aedeagus; b spur, basal spur; cgonst, clasper of gonostylus; db clasper, dorsal branch of clasper of gonostylus; db lobe, dorsal branch of lobe of gonostylus; f, foramen of male tergite X; goncx, gonocoxite; la spur, latero-apical spur; lb sh, lateral branch of sheath of aedeagus; lgonst, lobe of gonostylus; lh, lateral humps; mh, medial humps; pb sh, posterior branches of sheath of aedeagus; t9, male tergite IX; t10, male tergite X; vb clasper, ventral branch of clasper of gonostylus; vb lobe, ventral branch of lobe of gonostylus.   Figure 2.Schematic illustration of the male terminalia of representatives of  Amphineurus(dorsal view). A A. (A.) maculosus; B A. (Nothormosia) edentulus; C A. (A.) perdecorus; D A. (Nesormosia) fatuus. - Abbreviations: aed, aedeagus; aed sh, sheath of aedeagus; cgonst, clasper of gonostylus; db clasper, dorsal branch of clasper of gonostylus; goncx, gonocoxite; gonst, gonostylus; lb sh, lateral branch of sheath of aedeagus; lgonst, lobe of gonostylus; ms lobe, mesal lobe of gonocoxite; t9, male tergite IX; vb clasper, ventral branch of clasper of gonostylus.  Alexander (1920)allocated  Amphineurusas a subgenus of  Ormosiaafter highlighting the abundant and conspicuous wing macrotrichia (Fig. 5B). Edwards (1923)redefined the genus to support two New Zealand species. Edwards'diagnosis includes the short mid-leg, short and hairy wing, and radius four-branched. Nevertheless, these features occur in other Chioneinae. Edwards (1923)also referred to the fact that the medial vein forks in M3+4 and M1+2 and that the anal vein is sinuous. These features are plastic in the genus. Alexander (1929a)used the definition of Edwards (1923)and emphasized the characteristic "abundant trichiation" of the genus. However, in the same article, Alexander (1929a)described the genus  Maietta, which has more wing vestiture than  Amphineurus. Some species of the Neotropical subgenus A. (Rhamphoneurus)have less macrotrichia than other subgenera. No more inclusive diagnosis was proposed and recent revisions (e.g., Theischinger 1994, 1996) redefined only the diagnosis of the subgenus A. (Amphineurus).   Amphineurushas an 180 °rotation of male terminalia (in the words of Edwards: "undergone permanent torsion"). Edwards (1923)also shows that  Amphineurushas a long, finger-like gonocoxite process (Fig. 1A). This is evidence of the bifid gonocoxite, a condition present (Fig. 29G) in some Chioneinaewith inverted male terminalia. This gonocoxal process forms a protective hood ( Kramer 2020) for the appendices of the inverted terminalia. The Chioneinaegenera with inverted terminalia bear great diversity and frequently they have hairy wings similar to  Amphineurus. The great amount of wing vestiture resembles  MaiettaAlexander. The wings of some A. (Amphineurus)(Fig. 32: Node 18) have a distinctive radial sector such as seen in  MolophilusCurtis. Some A. (Amphineurus)(Fig. 32: Node 23) resembles  Hoplolabis(  ParilisiaSavchenko). The trifid claspers of A. (Rhamphoneurus)are observed in genera such as  HoplolabisOsten Sacken and  CheilotrichiaRossi. Despite that, in the group of genera with a bifid gonocoxite and inverted terminalia, vein R2+3 is normally aligned with R2+3+4. The exception is a clade of A. (Amphineurus)(Fig. 32: Node 18) that shows resemblances in the radial sector with  Molophilus(where R2+3+4 is lacking), but the former has a closed discal cell among other wing features which places it within  Amphineurus.