Remarkable New Genus of Gumillinae (Neuroptera: Osmylidae) From the Jurassic of China

SYSTEMATICS Remarkable New Genus of Gumillinae (Neuroptera: Osmylidae) From the Jurassic of China QIANG YANG, 1 VLADIMIR N. MAKARKIN, 1,2,3 AND DONG REN 1 Ann. Entomol. Soc. Am. 103(6): 855Ð859 (2010); DOI: 10.1603/AN10097 ABSTRACT Allotriosmylus uniramosus n. gen., n. sp. (Osmylidae: Gumillinae) is described from the Jurassic of China. This is the smallest representative of the family and possesses several peculiar features not occurring in other osmylids, such as a structurally well-formed clavus in the hind wing and a single-branched Rs in both the fore- and hind wings. KEY WORDS Neuroptera, Osmylidae, Gumillinae, Jurassic, Daohugou 1 College of Life Science, Capital Normal University, Beijing, 100048, China. 2 Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, 690022, Russia. 3 Corresponding author, e-mail: vnmakarkin@mail.ru. The extant Osmylidae is a relatively small neuropteran family with 200 known species distributed in warmtemperate to tropical regions of the Old Word and South America but that is surprisingly absent from North America (Oswald 1994, 2007). It is represented by many fossil genera going back to the Early Jurassic (Ansorge 1996). Mesozoic osmylids were diverse and are abundant in some localities; they are especially numerous at the Chinese locality of Daohugou, where approximately half of the 2,000 specimens examined were found to belong to this family (personal data). The Gumillinae is one of the most interesting subfamilies of the Osmylidae. In this group, the antennae are very long (much longer than their forewings), they possess an enlarged scapus, and their wings bear various venational peculiarities (Menon and Makarkin 2008). Only one extant genus, Gumilla Navás (one to two species from Brazil), and Þve Mesozoic genera (seven species) have been previously placed in the subfamily: Epiosmylus panfilovi Ren et Yin, Enodinympha translucida Ren et Engel, Nilionympha pulchella Ren et Engel, Nilionympha imperfecta Ren et Engel, Tenuosmylus brevineurus Wang et al. 2009 (all from the Middle/Late Jurassic of Daohugou, China); Epiosmylus longicornis PanÞlov (Late Jurassic of Karatau, Kazakhstan) and Nuddsia longiantennata Menon et Makarkin (Early Cretaceous [Late Aptian] of Crato Formation, Brazil) (PanÞlov 1980, Ren and Yin 2002, Ren and Engel 2007, Menon and Makarkin 2008, Wang et al. 2009). In addition, a species described by Ponomarenko (2003) as Nymphites cf. lithographicus from the Late Jurassic of Solnhofen (Germany) in Nymphitidae might be a gumilline (Menon and Makarkin 2008). Epiosmylus is the type genus of the extinct family Epiosmylidae, described by PanÞlov (1980). This family was considered closely related to, or a synonym of, Osmylidae (Makarkin 1990a, Ponomarenko 1990, Makarkin and Archibald 2003). Now, it is either treated as a subfamily of Osmylidae (Ren and Engel 2007; Engel and Grimaldi 2008) or synonymized with the Gumillinae (Menon and Makarkin 2008, Wang et al. 2009). Here, we describe a new genus and species of gumilline osmylid Allotriosmylus uniramosus n. gen. n. sp. from the Middle/Late Jurassic locality of Daohugou, closely related to Epiosmylus. This new genus is remarkable in particular for the presence of a structurally well-formed clavus in the hind wings, and the single branch of Rs in both the fore- and hind wings. Materials and Methods This study is based on one specimen collected near Daohugou Village (Shantou Township, Ningcheng County, Inner Mongolia, China) and housed in the fossil insect collection of the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Science, Capital Normal University, Beijing, China (CNUB; D.R., curator). These insect-bearing beds are here considered as belonging to the Jiulongshan Formation and dated Middle Jurassic (alternatively considered as belonging to the Daohugou Formation and dated latest Middle Jurassic to earliest Late Jurassic by Zhang 2010). Specimens were examined using an MZ12.5 dissecting microscope (Leica, Wetzlar, Germany), illustrated with the aid of Photoshop CS3 (Adobe Systems, Mountain View, CA); photographed with an SMZ1000 stereomicroscope (Nikon, Tokyo, Japan). We use the traditional (sensu Wootton 2003) venational terminology of Comstock (1918) following the recent interpretations of Oswald (1993), Archibald and Makarkin (2006), and Wedmann and Makarkin (2007). 0013-8746/10/0855Ð0859$04.00/0 2010 Entomological Society of America

856 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 6 Fig. 1. A. uniramosus n. gen., n. sp., photograph of the holotype CNU-NEU-NN2009700. Scale bar 1 mm. Systematic Palaeontology Order Neuroptera Linnaeus, 1758 Family Osmylidae Leach, 1815 Subfamily Gumillinae Navás, 1912 Genus Allotriosmylus n. gen. Type Species. Allotriosmylus uniramosus n. sp. Etymology. Allotri(o)- (from Greek allotrios, another kind) Ð(o)smylus (from Osmylus, a genus-group name), in references to the venation unlike that of other osmylids. Gender masculine. Diagnosis. May be easily distinguished from other genera of Gumillinae by only one branch of Rs in both wings (more than four in others), or by only one branch of CuA in the hind wing (more than three in others). Species Included. Type species only. Allotriosmylus uniramosus n. sp (Figs. 1 and 2AÐD) Holotype. CNU-NEU-NN2009700 (part), Capital Normal University, Beijing, China. A very incomplete, rather well-preserved, crumpled specimen: fragmentary thorax (probably in ventral aspect) with one pair of nearly complete wings, another pair only proximally preserved, and poorly preserved legs. Daohugou locality, Inner Mongolia, China; Middle Jurassic (Jiulongshan Formation). Etymology. The speciþc epithet is derived from the Latin unus, one, and ramosus, branched, in references to the single branch of Rs in each wing. Diagnosis. As for genus. Description. Foreleg(?) femur, tibia rather short; basitarsus longer than either of four other tarsomeres; at least femur, tibia and basitarsus covered with dense short hairs. Forewing elongate ovoid with rather rounded apex, 10.9 mm long, 3.4 mm wide. Trichosors poorly developed apically; possibly, two or three trichosors between each vein at apical posterior wing margin (poorly visible). Costal space relatively narrow, only slightly dilated at one Þfths its length. Subcostal veinlets simple, straight, rather widely spaced, 23 in number; postpterostigmal veinlets of Sc R1 simple. Pterostigma distinct, elongate, fuscous; incorporated veinlets rather clearly visible. Sc, R1 stout, fused at beginning of pterostigma. Sc R1 entering wing margin before apex. Subcostal space narrow, with one basal crossvein. Rs originates far from wing base, strongly zigzagged, with one long zigzagged branch. R1 space (between R1, Rs) broad, enlarged particularly at origin of Rs branch, with rather regularly arranged crossveins; one additional (probably aberrant) crossvein between those of R1 space. M appears fused basally with R for a rather long distance, dividing into MA and MP far distal from origin of Rs. Short oblique basal crossvein r-m near apparent origin of M. MA, MP diverging to nearly mid-point of wing, then slightly convergent; as a result, medial space very broad. MA nearly straight for most of length with shallow terminal fork, pectinately branched apically with four rather closely spaced branches. MP zigzagged, with seven widely spaced pectinate branches, proximalmost deeply forked. MP, CuA strongly approach proximally. Cu dividing into CuA, CuP very near to wing base. CuA relatively short, straight, with two apical pectinate branches. CuP very thin basally, somewhat stouter distally; pectinate with two branches. CuP, 1A

November 2010 YANG ET AL.: GUMILIINAE FROM JURASSIC OF CHINA 857 Fig. 2. Wing venation of A. uniramosus n. gen., n. sp., holotype CNU-NEU-NN2009700. (A and B) Forewings. (C and D) Hind wings. Scale bar 1 mm. Abbreviations: 1AÐ3A, Þrst to third anal veins; CuA, anterior cubitus; CuP, posterior cubitus; MA and MP, anterior and posterior branches of media (M); R1, Þrst branch of radius; Rs, radial sector; Sc, subcosta. strongly approached proximally. 1A long, with one long distal branch. 2A arched, simple (right wing) or with deep fork (left wing). 3A pectinate, with two branches. Between anal veins there are two to three crossveins. Anal region forms rather well developed clavus; claval furrow not detected. Hairs along wing margin rather short, somewhat longer along basal portion of wing and on longitudinal veins, much longer on longitudinal veins in proximal portion of wing. Jugal lobe absent or not preserved. Hind wing elongate, with acute apex, 9.8 mm long, 2.7 mm wide. Apical trichosors poorly-developed. Costal space narrow, slightly dilated medially. Subcostal veinlets simple, straight, widely spaced, 17 in number; veinlets of Sc R1 distal to pterostigma simple, oblique. Sc, R1 fused at beginning of pterostigma; Sc R1 enters margin before wing apex. Pterostigma distinct, fuscous, elongate; incorporated veinlets poorly visible, some appear strongly oblique. Sc, R stout for entire length; considerably thinner after fusion. Subcostal space very narrow; crossveins not detected. Rs zigzagged, with one branch. R1 space (between R1, Rs) similar to that of forewing. Basal sinuate crossvein m-r absent. M fused basally with R for a rather long distance, dividing into MA, MP near to wing base. Medial space very broad. MA straight for most length, almost not zigzagged, entering margin after wing mid-point; distal portion not preserved. MP long, zigzagged in distal portion, with six preserved simple branches. MP, CuA strongly approach proximally. Cu dividing into CuA, CuP very near to wing base. CuA short, with one long simple branch; CuP short, unforked; two crossveins between CuA and CuP. 1A, 2A and 3A short, simple; one crossvein between 1A, 2A, one between 2A, 3A. Anal region covers restricted area, represents very structurally well formed clavus (clearly exserted; alternatively, hind margin clearly emarginated between clavus, remigium); claval furrow not detected. Trichiation as in forewing but appears less dense on longitudinal veins. Jugal lobe absent or not preserved. Discussion The wing venation of Allotriosmylus gen. nov. is unusual. It superþcially resembles that of some chrysopoid families (here considered as Mesochrysopidae, Chrysopidae and Ascalochrysidae: Ren and Makarkin 2009), especially Mesochrysopidae by its reticulation and apically fused Sc and R1; however, more detailed analysis excludes it from the superfamily. It differs signiþcantly from Mesochrysopidae by the presence of trichosors, the elongate M (which does not exceed mid-wing in Mesochrysopidae) in the forewing, and

858 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 103, no. 6 the presence of CuP in the hind wing (lost in Mesochrysopidae). Other chrysopoid families are more dissimilar. Allotriosmylus gen. nov. shares a number of venational character states with Epiosmylus PanÞlov, also known from Daohugou, suggesting that they are most closely related and belong together in the osmylid subfamily Gumillinae, even though Allotriosmylus lacks the very long antennae and enlarged scapus considered as diagnostic for the subfamily (see Menon and Makarkin 2008) present in the Epiosmylus type species, E. longicornis (PanÞlov 1980). The wings of Allotriosmylus gen. nov. differ from those of Epiosmylus by the single branch of Rs (four to six in Epiosmylus) in both wings; the single branch of 1A (three in Epiosmylus); the greatly dilated medial space (slightly dilated in Epiosmylus) in the forewing, and the welldeveloped clavus in the hind wing (see below). Other gumilline genera are more dissimilar and probably more distantly related. The structure of the anal region (i.e., the clavus) is most remarkable in this species. Its margin is clearly exserted in the hind wing; it can be interpreted also as the hind margin clearly emarginated between the clavus and the remigium. In the majority of other Neuroptera, the entire hind margin of the wing is usually smooth (except the species with clearly undulated hind margin, e.g., Palaeogetes ponomarenkoi Makarkin, 1990; Makarkin 1990b). Only in some genera is the clavus slightly exserted, for example, in the hind wings of Epiosmylus, Platystoechotus Carpenter (Polystoechotidae), Plega Navás (Mantispidae), Nevrorthus Costa and Nipponeurorthus Nakahara (Nevrorthidae), and in the forewing of Anchieta Navás (Mantispidae) (pers. data). The wing shape and the structure of the clavus (including its simple venation) of Allotriosmylus uniramosus quite resembles those seen in many Mecoptera wings and Plecoptera forewings (cf. Ennos and Wootton 1989: Þg. 1; Béthoux 2005: Þgs. 1Ð5). The clavus differs greatly between the forewing and the hind wing in this species. In the hind wing, it is well differentiated from the remigium, however, it covers a very restricted area with simple venation; this is reversed in the forewing, where it is less distinctly formed, yet covers a much larger area. This difference is found also in Panorpa Linnaeus (Mecoptera) whose wings have a similar shape (Ennos and Wootton 1989). These differences between the fore- and hind wings are associated with ßight performance, affecting their pitch control (Ennos and Wootton 1989; Wootton 1992). Another unusual character found in Allotriosmylus gen. nov. is the single-branched Rs. This condition is probably unique among medium-sized neuropterans, which, as a rule have a pectinately branched Rs. The only other known group with a single-branched Rs is Coniopterygidae, minute neuropterans whose wings are a maximum of 5 mm long and have strongly reduced venation. The single-branched Rs occurs also in a minute berothid (forewing 3.36 mm long) (Archibald and Makarkin 2004: Þg. 3). Having trichosors restricted to only the apical portions of the wings is a condition characteristic of the subfamily; in the majority of other osmylids these are more widely distributed on wing margins. In Allotriosmylus uniramosus, however, there seem to be several (up to three) trichosors between vein tips along the posterior margin in the apical portion of the wing. 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