Protoliupanshania wangi, Huang, Di-Ying & Nel, Andre, 2010

Huang, Di-Ying & Nel, Andre, 2010, Protoliupanshania wangi, a new genus and species from the Chinese Early Cretaceous (Odonata: Aeshnoptera: Liupanshaniidae), Zootaxa 2387, pp. 57-62 : 58-61

Protoliupanshania wangi gen. nov., sp. nov.

( Figs. 1–6 )

Derivation of name. Named for Mr. Wang Xiang-Dong, who collected and gave us the holotype.

Material. Holotype NIGP 148199, a print of a well-preserved complete hind wing; paratype specimen NIGP 148200, a poorly-preserved specimen with part and counterpart with head trunk and four wings. Types housed in Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, China.

Type locality. Lower Cretaceous Yixian Formation (ca. 125 Ma); Beipiao City, Liaoning Province, NE China.

Diagnosis. That of the genus.

Description. Holotype: print of a complete hind wing, 45.5 mm long, wing 16.1 mm wide; distance from base to arculus 5.0 mm; from arculus to N 15.1 mm; from arculus to base of RP 3/4 9.2 mm; from N to Pt 14.6 mm, from N to wing apex 25.7 mm, from Pt to wing apex 6.4 mm; Pt 5.8 mm long, 1.1 mm wide, covering five cells; pterostigmal brace oblique and aligned with basal side of Pt; no secondary antenodal cross-vein basal to Ax1, two antenodal cross-veins of first row and one of second row between Ax1 and Ax2, six of first row and six of second row distal of Ax2; distance from wing base to Ax1 3.9 mm, between Ax1 and Ax2 6.1 mm; Ax2 aligned with apex of T; RP and MA strongly separated at arculus; hypertriangle 5.3 mm long, free of cross-veins; T 1.8 mm distal of arculus, free of cross-veins, broad, with basal side 1.9 mm long, anterior side 3.3 mm long, and MAb 3.7 mm long; a distinct angle in MAb at base of convex tp; part of MAb distal to tp long, 2.8 mm; postdiscoidal area with two rows of cells; a strongly developed convex tp, long and slightly zigzagged; no Mspl; basal part of area between RA and RP with six cross-veins basal to RP 3/4, and five distal of base of RP 3/4 and basal of subnodus; area between RP and MAa with five cross-veins basal to RP 3/4; one visible oblique vein ‘O’ three cells distal to base of RP 2; one row of cells between IR2 and RP 3/4 two cells distal to subnodus; MAa and RP 3/4 parallel, nearly straight; one row of cells between RP 3/4 and MAa basal of level of vein ‘O’, two rows distally; one row of cells between MP and CuAa, with no narrowing of area between these veins opposite base of RP 3/4; median and submedian area, and subdiscoidal space free; CuP and PsA strongly curved; posterior wing margin rounded at base (indicating a female specimen); anal area long and broad, with three posterior branches of AA directed towards posterior wing margin, and 9-10 rows of cells between AA and posterior wing margin; al small, opened posteriorly with no strong cross-vein between AA and CuAb, composed of five cells; cubito-anal area broad, with seven rows of cells between CuAa and posterior wing margin; CuAa with six posterior branches; no Rspl, but four posterior longitudinal secondary veins between IR2 and posterior wing margin; base of RP 2 aligned with subnodus; no well-defined IR1; one row of cells between RP 1 and RP 2 between subnodus and two cells basal of Pt, two rows distally.

Paratype: part and counterpart of a body with the four wings in connection. Wings and body are strongly deformed and their dimensions cannot be measured with accuracy. Eyes separated; abdomen with long cerci. All preserved structures of hind wings identical to those of holotype hind wing, including absence of Mspl and Rspl, al posteriorly opened, shape of T and tp, etc. Fore wing venational details largely undecipherable or not visible: hypertriangle longer than T, free of cross-veins; T free of cross-veins, broad, equilateral; median, submedian, and subdiscoidal spaces free of cross-veins; PsA strong; postdiscoidal area with four rows of cells; a strong convex tp, long and slightly zigzagged; no Mspl; no Rspl; Pt elongate, covering four cells, pterostigmal brace oblique and aligned with basal side of Pt; two rows of cells in anal area; four rows of cells in cubito-anal area; at least one secondary antenodal cross-vein between Ax1 and Ax2; Ax2 nearly opposite distal side of T.

Discussion. Protoliupanshania wangi shares with Liupanshaniidae Bechly et al., 2001 the following characters: both pairs of wings with a strong convex secondary longitudinal vein (tp) in the postdiscoidal area originating from a strongly angulate MAb (convergent to several other groups of Aeshnoptera and to the clade Gomphides Bechly et al., 1998); unique shape of elongate and narrow hind wing T (anterior side of T distally curved and ending on anterior side MA of hypertriangle); forewing T broad and nearly equilateral; distal accessory oblique vein between RP 2 and IR2 absent.

Similar shape tp and T also occur in the petalurid family Cretapetaluridae Nel et al., 1998, but Cretapetalura Nel et al., 1998 has its pterostigmal brace strongly recessed basally ( Nel et al., 1998). Also the aeshnopteran Va l d a e s h n a Jarzembowski, 1988 and the gomphid Hageniidae Tillyard & Fraser, 1940 and Lindeniidae Jacobson & Bianchi, 1905 (sensu Bechly 1991) have tp, but their T of both fore- and hind wings have the same elongate shape, unlike Protoliupanshania . Enigmaeshna Nel et al., 2008 (Enigmaeshnidae, only hind wing known) has a strong tp but it differs from Protoliupanshania by the longer anal area with a posteriorly closed al, and hypertriangle, T and subdiscoidal space crossed. Lin et al. (2004: Fig. 2) figured a zigzagged tp for the Chinese Cretaceous Fuxiaeschna hsuifunia Lin et al., 2004 which they included in the aeshnopteran family Rudiaeschnidae Bechly et al., 2001 , but this character is not shared by the type genus Rudiaeschna Ren & Guo, 1996. Fuxiaeschna differs from Protoliupanshania by the presence of a closed al and broader hind wing T. Lastly the representatives of the clade Neoaeshnida Bechly, 1996 ( Gomphaeschnidae + Aeshnodea) have developed tp but they have well defined Rspl and Mpsl, and a posteriorly closed hind wing al unlike Protoliupanshania .

Other liupanshaniid synapomorphies are less pronounced or absent in Protoliupanshania ; its hind wing MAb is sigmoidally curved, but distinctly less so than in other Liupanshaniidae. The hind wing T of Protoliupanshania is free instead of being divided into several cells by parallel cross-veins; its forewing T is free instead of being divided into cells (forewing T only known in Araripeliupanshania Bechly et al., 2001 and Paramesuropetala Bechly et al., 2001). Also the subdiscoidal space of Protoliupanshania is larger and broader than in other Liupanshaniidae ( Bechly et al., 2001; Lin et al., 2002).

Protoliupanshania can be considered a member of the family Liupanshaniidae, but it could be the sister genus of all other taxa in this family, because of the plesiomorphic character state ‘tp basally zigzagged’ (hind wing tp not basally zigzagged in Araripeliupanshania Bechly et al., 2001, Guyuanaeschnidia Lin, 1982, Liupanshania Hong, 1982 , and Paraliupanshania Bechly et al., 2001); but some uncertainty remains due to the uncertain phylogenetic position of Paramesuropetala Bechly et al., 2001. If the basal position of Protoliupanshania is confirmed after a better knowledge of Paramesuropetala, this might suggest Central Asia as an origin for this group. The absence of Rspl and Mspl in Protoliupanshania is also found in Guyuanaeschidia Lin, 1982, Liupanshania Hong, 1982 , while Paraliupanshania Bechly et al., 2001 have well defined Rspl and Mspl.

Figures

FIGURE 1. Protoliupanshania wangi gen. nov., sp. nov., holotype NIGP 148199, photograph (scale bar 5 mm). FIGURE 2. Protoliupanshania wangi gen. nov., sp. nov., paratype specimen NIGP 148200, photograph (scale bar 5 mm).

FIGURES 3 – 6. Protoliupanshania wangi gen. nov., sp. nov., 3. holotype NIGP 148199, drawing. 4 – 6. Paratype specimen NIGP 148200. 4, left hind wing. 5, left forewing. 6, right forewing, standard abbreviations: AA anal vein, Ax 1 Ax 2 primary antenodal cross-veins, CuAa distal branch of cubitus anterior, CuAb proximal branch of cubitus anterior, IRi intercalary radial veins, MAa distal branch of median anterior, MAb posterior branch of median anterior, MP median posterior, N nodus, ‘ O’ lestine oblique vein, PsA pseudo-anal vein, Pt pterostigma, RA radius anterior, RP radius posterior, tp trigonal planate, T discoidal triangle, al anal loop (scale bars 5 mm).