A new campterophlebiid damsel-dragonfly (Odonata: Isophlebioidea) from the Middle Jurassic Yanan Formation of Yulin City, Shaanxi Province, NW China

The family Campterophlebiidae Handlirsch, 1920 is the dominant Jurassic clade of Odonata Fab-ricius, 1793, especially hosting a high diversification in northern China. The Chinese campter-ophlebiid damsel-dragonflies were mainly recovered from the Middle Jurassic of Inner Mongolia, northern China. In the present study, a new campterophlebiid, Parasinitsia qingyunensis n. gen., n. sp., is described from the early Middle Jurassic Yanan Formation of the Ordos Basin, NW China.


Comptes Rendus Palevol est indexé dans / Comptes Rendus Palevol is indexed by:
-Cambridge Scientific Abstracts -Current Contents® Physical -Chemical, and Earth Sciences ® -ISI Alerting Services ® -Geoabstracts, Geobase, Georef, Inspec, Pascal -Science Citation Index ® , Science Citation Index Expanded ® -Scopus ® .  Fleck & Nel 2002;Nel et al. 2008Nel et al. , 2009Zheng et al. 2016Zheng et al. , 2017Zheng et al. , 2018Zheng et al. , 2019Huang et al. 2018 The specimen was examined dry using a Nikon SMZ1000 stereomicroscope. Observation was augmented by temporary wetting with laboratory alcohol which improved the contrast between the fossil and the matrix, eliminating the surface irregularity of the latter. Photographs were taken using a Canon 5D digital camera and the line drawings were prepared from photographs using image-editing software (CorelDraw X7 and Adobe Photoshop CS6). The specimen is housed in the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS). All taxonomic acts established in the present work have been registered in ZooBank.
The higher classification of fossil and extant Odonatoptera is based on the phylogenetic system of Bechly (1996). The nomenclature of the dragonfly wing venation used in this paper is based on the interpretations of Riek (1976) and Riek & Kukalová-Peck (1984), as modified by Nel et al. (1993) and Bechly (1996). diAgnosis. -One row of cells in basal areas between MAa and MP, and between MP and CuAa; width of basal areas between MAa and MP, and between MP and CuAa nearly identical; CuAa approaching MP, with at least three rows of cells between it and posterior wing margin distally; area between MAa and MP widened before N, with at least three rows of cells in broadest area; MP with many posterior intercalary veins, with 15 rows of cells between it and posterior wing margin in broadest area; 'O' seven cells distal of Sn; a strong and long oblique vein between IR1 and RP2, oriented towards the wing base, lying slightly close to level of Pt base than to IR1 base; Pt long and elongate, covering six cells and not braced; two or three rows of cells between RA and RP distal of Pt. diAgnosis. -The same as for the monotypic genus.

AbbreviAtions
mAteriAl. -Holotype, NIGP163538a, b, part and counterpart of a well-preserved forewing, with posterior wing base and middledistal wing half preserved (Fig. 2); deposited in NIGPAS.  Area between MAa and RP3/4 widened distally, with many cells along below Pt. Area between RP3/4 and IR2 with one row of cells basal of N level, expanded distally with six rows of cells below N. Area between IR2 and RP2 with one row of cells slightly before Pt, expanded distally with 2-4 rows of cell below Pt. Area between RP2 and IR1 with one rows of cells slightly basal of Pt base, widened distally with many rows of cells. Area between IR1 and RP1 with two rows of cells before Pt, widened distally with many rows of cells. Area between RA and RP1 before Pt end, widened distally with two or three rows of cells.

DISCUSSION
Parasinitsia qingyunensis n. gen., n. sp. shares the typical characters of Campterophlebiidae: a medium size, no secondary antenodal crossveins between C and ScP, and basally opened discoidal cell in the forewing (Nel et al. 2009;Li et al. 2012a). Parasinitsia qingyunensis n. gen., n. sp. has similar widths of the basal parts of the postdiscoidal area between MAa and MP, and that between MP and CuA, and only one row of cells between these veins in their basal parts, only shared by the following campterophlebiid damsel-dragonflies: Ctenogampsophlebia Petrulevičius, Huang & Nel, 2011, Honghea Zheng, Shi, Wang, Chang, Dou, Wang & Zhang, 2017, Karatawia Martynov, 1925, Melanohypsa Pritykina, 1968, Parazygokaratawia Huang, Cai & Nel, 2018, Sinitsia, and Zygokaratawia Nel, Huang & Lin, 2008. Ctenogampsophlebia and Honghea have smaller wing sizes, MAa ending on MP, and fewer rows of cells between the main veins below the pterostigma, showing differences with Parasinitsia qingyunensis n. gen., n. sp. (Petrulevičius et al. 2011;Zheng et al. 2017). Karatawia has MAa zigzagged distally, one row of cells between RA and RP1 distal of the pterostigma, MP with fewer rows of cells between it and the posterior wing margin, differing from P. qingyunensis n. gen., n. sp. (Nel et al. 1993;Li et al. 2012b). Melanohypsa Pritykina, 1968, although only preserved with wing base, is similar with P. qingyunensis n. gen., n. sp. in sharing with an acute distal angle of the discoidal cell and two rows of cells in the anal area; it, however, differs from P. qingyunensis n. gen., n. sp. in having CuAa remaining parallel with MP instead of making a strong anterior curvature towards it (Pritykina 1968;Nel et al. 1993). Parazygokaratawia and Zygokaratawia have smaller sizes (30.8 and 35.5 mm, respectively), MAa zigzagged and weakened distally, a short pterostigma covering two or three cells, and less rows of cells between the main veins below the pterostigma, excluding any affinities with P. qingyunensis n. gen., n. sp. (Nel et al. 2008;Huang et al. 2018). Sinitsia quite resembles P. qingyunensis n. gen., n. sp. in having CuAa moving towards MP distally, no constriction between RP2 and IR2, many rows of cell between the main veins in the distal half of wing, plus a strong and long oblique vein between IR1 and RP2, oriented towards the wing base, and lying slightly close to the level of the pterostigmal base than to IR1 base, and a long pterostigma (Pritykina 2006;Nel et al. 2009). Parasinitsia qingyunensis n. gen., n. sp. differs from Sinitsia in having the oblique vein 'O' seven cells distal of Sn, two or three rows of cells between RA and RP1 distal of the pterostigma, at least three rows of cells in the broad part of postdiscoidal area basad the nodus, up to eight rows of cells in the area between IR2 and RP3/4 below the pterostigma, vs only four in Sinitsia, and c. 15 rows of cells between MP and posterior wing margin in the broadest area, vs 13 in Sinitsia (Pritykina 2006;Nel et al. 2009). Some Campterophlebiidae genera are based on incomplete wings in which the basal part of the postdiscoidal area is not preserved, e.g. Sarytashia Pritykina, 1970. This genus differs from the new fossil in the areas between IR2 and RP2 and between RP2 and IR1 much narrower (Nel et al. 1993).
Within Campterophlebiidae, Parasinitsia qingyunensis n. gen., n. sp. has many rows of cells between the main veins in the distal part (especially for the present of two or three rows of cells between RA and RP1 distal of the pterostigma), area between MAa and MP broadened in the middle part and distally constricted, many secondary intercalary veins originating from MP, 'O' seven cells distal of RP2 base, a strong and long oblique vein between IR1 and RP2 oriented toward the wing base, lying slightly close to the level of Pt base than to IR1 base and Pt covering many cells, all characters shared by Parafleckium Li, Nel, Ren & Pang, 2012(Li et al. 2012a. Parafleckium however, obviously differs from Parasinitsia qingyunensis n. gen., n. sp. in having a broad area between MP and CuAa, which is three times the size of the postdiscoidal area, MP with fifteen rows of cells between it and the posterior wing margin in the broadest part, and CuAa with only one row of cells between it and the posterior wing margin. Junfengi yulinensis Zheng & Zhang, 2017 is based on a male hind wing discovered from the Yanan Formation of Yulin (Zheng et al. 2017). Any affinities between Junfengi Zheng & Zhang, 2017 and P. qingyunensis n. gen., n. sp. can be excluded, since the former has MAa nearly parallel with MP, MP with fewer rows of cells between it and the posterior margin, the oblique vein 'O' three cells distal of Sn, fewer postnodal crossveins before the pterostigma, and IR2 with only one row of cells between it and RP3/4 before the level of IR1.
In conclusion, Parasinitsia qingyunensis n. gen., n. sp. resembles the campterophlebiid dragonfly Sinitsia and Parafleckium, but still strongly differs from these genera. We therefore justify a new genus for this unique damsel-dragonfly.

CONCLUSIONS
A new Campterophlebiidae, Parasinitsia qingyunensis n. gen., n. sp., is described from the Middle Jurassic Yanan Formation in northern Ordos Basin, northwestern China. Parasinitsia n. gen. exhibits most similarities with Sinitsia, but distinguished from all other campterophlebiid damsel-dragonflies in having MP with fifteen rows of cells between it and the posterior wing margin, two or three rows of cells between RA and RP1 distal of the pterostigma. Sinitsia is only recorded from the Upper Jurassic in Translaikalia. The present discovery suggests