PROTOSMYLINAE Krüger, 1913a

Winterton, Shaun L., Martins, Caleb Califre, Makarkin, Vladimir, Ardila-Camacho, Adrian & Wang, Yongjie, 2019, Lance lacewings of the world (Neuroptera: Archeosmylidae, Osmylidae, Saucrosmylidae): review of living and fossil genera, Zootaxa 4581 (1), pp. 1-99 : 44-49

PROTOSMYLINAE Krüger, 1913a

Type genus. Protosmylus Krüger, 1913a: 30 .

Diagnosis. Ocelli present or variously reduced (medial ocellus sometimes reduced or absent), antennae shorter than forewing length; prothorax width subequal to length; pronotum length subequal to width; female forecoxae without processes (except Protosmylus ); wings ovoid; costal area broad basally, subcostal veinlets simple, sometimes irregularly forked in FW; trichosors present along entire wing margin; single FW sc-r crossvein basally; both wings with RP with ca. 8-10 branches (rarely more), rarely sinuous distally, radial area with relatively few crossveins basally resulting in large and well defined cells; two (rarely one or three) gradate series present and well defined, often distant from the other irregular radial crossveins in basal half of wing; FW M vein forked near wing base, near origin of RP1; CuA usually with more branches in HW than in FW; HW CuP simple and relatively short, not pectinately branched; male with abdominal tergites 8 and 9 as separate sclerites with eversible scent glands absent; male genitalia less sclerotised with gonarcus narrow, arched and not observable externally; parameres present, fused medially and arched; anterolateral arm of gonarcus non-articulating; female genitalia with gonopophysis 9 basally associated with gonocoxite 9 (not acting upon sternite 8); sternite 8 as small knob-like process located near anteroventral margin of tergite 9; spermathecae oblong and folded upon itself, spermathecae duct extremely long and coiled.

Comments. Protosmylinae comprises four extant and six extinct genera. Three of the extant genera are found in the Oriental and Eastern Palaearctic regions while the fourth genus is known only from a relatively small area in Ecuador. The six extinct genera are widely known from deposits throughout the Palaearctic and one in the Nearctic, ranging in age from the Middle Jurassic to late Eocene. Based on a phylogenetic analysis Lysmus was recently placed in Protosmylinae by Winterton et al. (2017). This subfamily has attracted attention recently, with revisions by Wang et al. (2010) ( Juraheterosmylus ), Dong et al. (2016) ( Heterosmylus ) and Winterton & Wang (2016) ( Gryposmylus ) have resulted in new species being described in these genera. Wang et al. (2010) revised the diagnosis of Protosmylinae , updating the original concept by Krüger (1913). Protosmylinae is closely related to Spilosmylinae ( Winterton et al., 2017a,b) and the two subfamilies share features such as a single to few branched hind wing vein CuP, narrow gonarcus and fused parameres in the male terminalia, as well as sternite 8 migrated posteriorly into a small know-like process in the female terminalia ( Winterton et al., 2017). The similarities between the two subfamilies are indeed striking and few reliable characters are known that separate them ( Krüger, 1913; Wang et al., 2010; Winterton et al., 2017) resulting to problems for placing taxa (e.g., Lysmus ), particularly for some fossils where only wing fragments are known. Within Protosmylinae , besides similarities in wing venation, male and female genitalic morphology is also relatively uniform, with only slight differences amongst genera and even some species. The distinction of various extant and extinct genera is reliant on minor wing venation differences that should be evaluated further in a comparative context to determine synonymies with the nominal genus Protosmylus (see comments below under Protosmylus ). Important characters useful for differentiating genera in this subfamily include the number of distinct gradate series, and the relative sizes and number of branches of CuP and CuA in the forewing ( Table 2 ). Larvae are unknown for this subfamily.

Genera included. Gryposmylus Krüger , Heterosmylus Krüger , Juraheterosmylus Wang et al. , Jurosmylus Makarkin & Archibald , Lysmus Navás , Mesosmylidus Jepson et al. , Osmylidia Cockerell , Paryphosmylus Krüger , Petrushevskia Martynova , Protosmylina Jepson et al. , Protosmylus Krüger , Pseudosmylidia Makarkin.

Tables

TABLE 2. Selected informative morphological characteristics of genera of Protosmylinae.

	Number and orientation of gradate series	Relative areas of FW CuA and CuP branching	Position of FW M fork
Gryposmylus	Two, divergent	CuA area smaller than CuP	Approx. level with RP1 fork
Heterosmylus	Three, subparallel	Subequal in area	Basal to RP1 fork
Juraheterosmylus	Three, subparallel	Subequal in area	Approx. level with RP1 fork
Jurosmylus	Three, subparallel	CuA area larger than CuP	Approx. level with RP1 fork
Lysmus	Two to three, subparallel	CuA area smaller than CuP	Approx. level with RP1 fork
Mesosmylidus	Unknown	CuA area smaller than CuP	Basal to RP1 fork
Osmylidia	Two, subparallel	Subequal in area	Approx. level with RP1 fork
Paryphosmylus	Three to four, subparallel	CuA area smaller than CuP	Approx. level with RP1 fork
Petrushevskia	One	CuA area smaller than CuP	Approx. level with RP1 fork
Protosmylina	Three, divergent	Unknown	Approx. level with RP2 fork
Protosmylus	Two, subparallel	Subequal in area	Approx. level with RP1 fork
Pseudosmylidia	Three, subparallel	CuA area larger than CuP	Approx. level with RP2 fork