Sinoagetopanorpidae fam. nov., a New Family of Scorpionflies (Insecta, Mecoptera) from the Guadalupian of South China

Simple Summary Mecopterans have been sparsely reported from the Permian of China, despite their great biodiversity in the fossil record in the world. Herein, we describe and illustrate three genera (two new genera) and eleven species (ten new species) belonging to a new family Sinoagetopanorpidae fam. nov. from the upper Guadalupian Yinping Formation of Anhui Province, China: Sinoagetopanorpa permiana Lin, Nel and Huang, 2010, S. nigra sp. nov., S. rotunda sp. nov., S. lini sp. nov., S. minuta sp. nov., S. elegans sp. nov., S. grimaldii sp. nov., S. magna sp. nov., Raragetopanorpa zhangi gen. et sp. nov., Permoagetopanorpa yinpingensis gen. et sp. nov. and P. incompleta sp. nov. Our new discovery indicates a high diversity of mecopterans in the Permian of China, and Signoagetopanorpidae might have evolved independently on the Yangtze Platform. Abstract Mecoptera was in great abundance in the Permian, but little is known from China. A new family, Sinoagetopanorpidae fam. nov., is described and illustrated from the upper Guadalupian Yinping Formation at Yinping Mountain, Chaohu City, Anhui Province, China. Sinoagetopaorpa permiana Lin, Nel and Huang, 2010 was previously attributed to Permochoristidae and now is revised as the type species of Sinoagetopanorpidae fam. nov. Three genera (two new genera) and ten new species of this new family are described and illustrated: Sinoagetopanorpa permiana Lin, Nel and Huang, 2010, S. nigra sp. nov., S. rotunda sp. nov., S. lini sp. nov., S. minuta sp. nov., S. elegans sp. nov., S. grimaldii sp. nov., S. magna sp. nov., Raragetopanorpa zhangi gen. et sp. nov., Permoagetopanorpa yinpingensis gen. et sp. nov. and P. incompleta sp. nov. Some isolated hind wings are described and illustrated, although it is difficult to assign them to any particular species. As a dominant mecopteran lineage in the Yinping Formation, Sinoagetopanorpidae represents an endemic group that might have independently evolved on the Yangtze Platform.


Introduction
The common name "scorpionflies" of the insect order Mecoptera is derived from the fact that some males bear unturned and bulbous genitalia that resemble the stingers of scorpions. Holometabolous insects have the richest biodiversity among all insect clades and can be dated back to the Pennsylvanian [1]. Mecoptera is one of the most ancient and morphologically generalized holometabolous insect orders, and they represent an important holometabolous group in the Permian. Mecoptera can be traced back to the beginning of the Permian [2] and thrived from the Permian to Cretaceous [3][4][5][6][7][8][9][10][11][12][13][14][15][16][17][18][19][20]. However, many mecopteran lineages went extinct during the Cretaceous, and a modern-looking mecopteran assemblage emerged in the Cenozoic. Extant mecopterans encompass ca. 700 species and 40 genera assigned to 9 families [21,22]. Bittacidae and Panorpidae are overwhelmingly diverse among mecopterans in the extant fauna. Mecoptera is more prosperous in the fossil record than it is in the Recent, with more than 700 species and 210 genera in approximately 40 families recorded to date [22][23][24].  Material. Twenty-four specimens, eleven specimens with part and counterpart, and ten complete or nearly so. Holotype, NIGP143428, wing base lacking a small part, with part and counterpart (Figures 2 and 3A,B); from the Yinping Formation, Chaohu City, China (Lin et al. [31], Figures 1-4).  Paratypes, NIGP200888 ( Figure 3C,D), a complete forewing, with part and counterpart; NIGP200889 ( Figure 3E,F), a forewing lacking a small part of apex, with part and counterpart; NIGP200890 ( Figure 3G,H), a complete forewing, with part and counterpart. Type locality and horizon. Yinping Mountain, Chaohu City, Anhui Province, China; Yinping Formation (Capitanian).
Material. Twenty-four specimens, eleven specimens with part and counterpart, and ten complete or nearly so. Holotype, NIGP143428, wing base lacking a small part, with part and counterpart (Figures 2 and 3A,B); from the Yinping Formation, Chaohu City, China (Lin et al. [31], Figures 1-4).  Description. Forewing length 5.6 mm, width 3.0 mm, W/L 1.9; apex rounded a base slightly shrunken; wing almost fully covered with dark color, with three small h line patches at costal area and seven small hyaline spots at wing base; Sc terminated at of wing, apical Sc3 extending with the same direction as basal stem Sc; R1 moderat curved apically; Rs five-branched, stem Rs1+2 length 1.2 mm, stem Rs3+4 length 0.3 m stem Rs4a+b shorter than its branches; a crossvein connecting Rs1+2 fork and Rs3; Rs merg with R1 at a distance of 1.5 mm from wing base; a crossvein connecting stem Rs3+4 a stem M1+2, Rs4b and M1, respectively; M with six branches, stem M1+2 as long as stem M both M2 and M4 bifurcated into two branches; stem M4 length 0.2 mm, a crossvein co necting stem M2 and M3, crossvein m-cua connecting basal M4b and CuA; CuA curv after the connection with corossvein m-cua, the crossvein between basal CuA and C oblique; CuP straight, apically curved; M + CuA fork proximal to R1 + Rs fork, R1 + Rs fo proximal to the first fork of Sc; two anal veins visible, a crossvein connecting A1 and near base.
Revised diagnosis. Forewing moderate in size, covered with dense dark-colored spots, a hyaline rounded triangular spot at apex of each interface between Rs and M branches.
Revised description. Forewing length 5.6-7.9 mm, width 2.9-4.0 mm, L/W 1.9-2.2; apex somewhat rounded; dark color lessening from wing apex toward wing base, a large patch of dark color at apical wing, basal wing with numerous dark-colored spots, costal area with 5-6 hyaline patches that devoid of dark color, a hyaline rounded triangular spot that devoid of dark color at apex of each interface between Rs and M branches; Sc terminated at about 4/5 of wing, with three evenly developed elongated branches; R 1 single, curved slightly apically; pterostigma large, with almost a half below the apical R 1 ; Rs five-branched, Rs 4 bifurcated into two branches, Rs 1 slightly curved upwards, Rs 1+2 fork does not reach the level of M 2a+b fork; a crossvein connected near Rs 1+2 fork and Rs 3 , stem Rs 3+4 and stem M 1+2 , Rs 4b and M 1 , respectively; M six-branched, M 2 bifurcated into two branches; the forking patterns of the three branches of M 3+4 variable; most specimens with M 3 single and M 4 two-branched, some with M 3 two-branched and M 4 single, and the other with the three branches of M 3+4 forking at one point; stem M 1+2 equal to or longer than stem M 3+4 , a crossvein connecting stems M 2 and M 3 , crossvein m-cua connecting stem M 4 (or posterior branch of M 4 ) and CuA; CuA curved apically, CuP single, straight; the crossvein between basal CuA and CuP oblique to horizontal; the holotypic specimen with R 1 + Rs fork proximal to the first fork of Sc and M + CuA fork, M + CuA fork slightly proximal to the first fork of Sc; most paratypic specimens with M + CuA fork proximal to R 1 + Rs fork and R 1 + Rs fork proximal to the first fork of Sc; two anal veins visible, a crossvein connecting each other near base.
Remarks. The holotype is slightly deformed, so the original line drawings (see Lin et al. [31]: Figures 3 and 4) are inaccurate. The upper branch of CuA in the original line drawing is in fact the lower branch of M 4 (M 4b ).
Etymology. The specific name is derived from the Latin word nigra, dark, referring to the dark-colored wing.
Diagnosis. Forewing almost fully with dark colors, absence of dark-colored spots, absence of a hyaline rounded triangular spot at apex of each interface between Rs and M branches; R 1+2 fork near the level of M 2a+b fork.
Description. Forewing length 5.6 mm, width 3.0 mm, W/L 1.9; apex rounded and base slightly shrunken; wing almost fully covered with dark color, with three small hyaline patches at costal area and seven small hyaline spots at wing base; Sc terminated at 2/3 of wing, apical Sc 3 extending with the same direction as basal stem Sc; R 1 moderately curved apically; Rs five-branched, stem Rs 1+2 length 1.2 mm, stem Rs 3+4 length 0.3 mm, stem Rs 4a+b shorter than its branches; a crossvein connecting Rs 1+2 fork and Rs 3 ; Rs merged with R 1 at a distance of 1.5 mm from wing base; a crossvein connecting stem Rs 3+4 and stem M 1+2 , Rs 4b and M 1 , respectively; M with six branches, stem M 1+2 as long as stem M 3+4 ; both M 2 and M 4 bifurcated into two branches; stem M 4 length 0.2 mm, a crossvein connecting stem M 2 and M 3 , crossvein m-cua connecting basal M 4b and CuA; CuA curved after the connection with corossvein m-cua, the crossvein between basal CuA and CuP oblique; CuP straight, apically curved; M + CuA fork proximal to R 1 + Rs fork, R 1 + Rs fork proximal to the first fork of Sc; two anal veins visible, a crossvein connecting A 1 and A 2 near base.
Sinoagetopanorpa rotunda sp. nov. ( Figure 5 and Figure 19C; ZooBank LSID urn:lsid:zoobank.org:act:9E2C0D32-5E2C-44E1-B69D-FDD9907493C6). Description. Forewing broad, broadest area at the 4/5 of wing, length 7.1 mm, width 4.1 mm, L/W 1.7; some elongated hyaline patches developed at anterior and basal wing, and a hyaline rounded triangular spot at apex of each interface between Rs and M branches; crossveins usually accompanied with a small hyaline patch; pterostigma large, with almost a half below the apical R1; Sc terminated at 2/3 of wing, with three evenly developed elongated branches; R1 single, curved distinctly in the pterostigma; Rs fivebranched, Rs1+2 fork proximal to M2a+b fork; stem Rs1+2 4 times as long as stem Rs3+4; Rs1 curved upwards in middle; stem Rs4a+b as long as its branches; a crossvein connecting basal Etymology. The specific name is derived from the Latin rotunda, rounded, indicating the rounded wing shape.
Diagnosis. Forewing distinctly oval-shaped, absence of dark-colored spots; a hyaline rounded triangular spot at apex of interface between each Rs and M branches; R 1 distinctly curved apically.
Description. Forewing broad, broadest area at the 4/5 of wing, length 7.1 mm, width 4.1 mm, L/W 1.7; some elongated hyaline patches developed at anterior and basal wing, and a hyaline rounded triangular spot at apex of each interface between Rs and M branches; crossveins usually accompanied with a small hyaline patch; pterostigma large, with almost a half below the apical R 1 ; Sc terminated at 2/3 of wing, with three evenly developed elongated branches; R 1 single, curved distinctly in the pterostigma; Rs five-branched, Rs 1+2 fork proximal to M 2a+b fork; stem Rs 1+2 4 times as long as stem Rs 3+4 ; Rs 1 curved upwards CuA and CuP simple, CuA curved after the crossvein m-cua; the crossvein between CuA and CuP curved; R 1 + Rs fork and M + CuA fork nearly at the same level and proximal to the first fork of Sc; A 1 and A 2 single, a crossvein connecting each other near base.
Paratypes. NIGP200914, a complete forewing, with part and counterpart ( Figure 5C,D,G), length 6.9 mm, width 4.0 mm, L/W ratio 1.7; lower margin in the middle concaved obviously; stem Rs 1+2 twice as long as stem Rs 3+4 ; stem M 1+2 slightly longer than M 3+4 and nearly twice as long as stem M 2a+b ; M 3 two-branched, M 4 single, stem M 3a+b short; crossvein m-cua connecting CuA with basal M 4 . NIGP200915 ( Figure 5E,F,H), a forewing with wing base not preserved, with part and counterpart, length 6.7 mm (as preserved), width 3.9 mm; stem Rs 1+2   Forewing relatively elongated, broadest at 2/3 of wing, right forewing length 7.5 mm, width 3.5 mm, L/W 2.1 (Figures 6E,F and 7C); three small hyaline patches located at costal area; R1 single and curved apically; Rs five-branched, a terminal fork visible at right forewing of holotype ( Figure 7C), left forewing ( Figure 7B) lacking the terminal fork; stem Rs1+2 about twice as long as stem Rs3+4, Rs1+2 fork proximal to M1+2 fork, a crossvein connecting basal Rs2 and Rs3, basal Rs4b and M1, stem Rs3+4 and stem M1+2, respectively; M sixbranched, stem M1+2 1.5 times as along as stem M3+4, both M2 and M4 two-branched; a crossvein connecting stem M2a+b and M3, crossvein m-cua connecting CuA with M near   Hind wing smaller than forewing; left hind wing length 6.4 mm, width 3.2 mm, L/W 2.0 (Figures 6G,H and 7D,E); wing with dense dark color, a large hyaline patch stretched from the middle part of wing base to middle wing, a hyaline rounded triangular spot at apex of interface between each Rs and M branches; costal area broad, Sc single, terminated at middle wing, curved apically; R1 bifurcated into two branches at middle wing; Rs fivebranched, stem Rs1+2 more than 3 times as long as stem Rs3+4; stem Rs4a+b longer than its branches, Rs1+2 forking at the same level as M1+2 fork; a crossvein between R1b and Rs1, basal Rs2 and Rs3, Rs3 and Rs4a, respectively, and the crossveins inside the small spots devoid of dark color; M five-branched, M2 two-branched, stem M1+2 as long as stem M3+4 and M2a+b; basal stem Rs4a+b and stem M1+2, Rs4b and M1, stem M2a+b and M3 connected by a crossvein, respectively; crossvein m-cua connecting M3+4 fork and CuA; CuA and CuP straight, the crossvein between basal CuA and CuP oblique; left forewing with R1 + Rs fork slightly proximal to M + CuA fork; right forewing with R1 + Rs fork distinctly proximal to M + CuA fork; two straight anal veins visible.
Paratypes: NIGP200917 (Figures 8 and 19D), forewing, length 6.4 mm, width 3.0 mm, W/L 2.1; three relatively large irregular hyaline patches covered at the area of costa, subcosta and along the R1, a hyaline rounded triangular spot at apex of each interface between Rs2-4 and M branches, some crossveins in spots and approximately 10 small spots at wing base; stem M1+2 slightly longer than stem M3+4 and twice as long as stem M2a+b; the three branches of M3+4 nearly forking at one point.
Remarks. The dark color inside the forewings of the holotype is too poorly preserved to reconstruct.
Etymology. The specific name is derived from the Latin minuta, small, indicating the small wing size. Diagnosis. Forewing small, a hyaline rounded triangular spot at end of each interface of Rs and M branches; Rs4 forking late, A3 short, the area between A3 and wing margin narrow.
Material. Only holotype (NIGP200920) examined, a nearly complete forewing, with part and counterpart (Figures 9 and 19E). Etymology. The specific name is in honor of the late paleoentomologist, Prof. Qibin Lin, for his extraordinary contribution to the paleoentomology of the Yinping Formation.
Diagnosis. Wing without dark-colored spots; a hyaline rounded triangular spot at apex of interface between each Rs and M branches; forewing with Rs 1+2 fork proximal to M 2a+b fork; hind wing devoid of dark color at wing base; R 1 bifurcating into two terminal branches.
Material. Four specimens, two with part and counterpart. Holotype, sex unknown, NIGP200916, with part and counterpart ( Figures 6 and 7), a specimen preserved with part of body and four wings, abdomen segments detected; left forewing and right hind wing overlap with each other, left forewing upturned. Paratypes, NIGP200917, a complete forewing, with part and counterpart (Figures 8 and 19D); other unillustrated specimens: NIGP200918 and NIGP200919.
Description. Body length 5.3 mm (as preserved), head and thorax preserved obscurely, thorax medium size; abdomen with seven clearly discernible segments, length shorter than width, last two segments narrowed; genitalia relatively small compared with abdomen, clip-shaped, basistyles and dististyles lacking details; the left middle leg preserved with femur, tibia and tarsus; femur robust, length 1.5 mm (as preserved); tibia length 1.9 mm; five tarsi segments preserved (with total length 1.5 mm); tarsomeres gradually shorten from base towards apex; two front legs and left hind leg partly preserved.
Forewing relatively elongated, broadest at 2/3 of wing, right forewing length 7.5 mm, width 3.5 mm, L/W 2.1 ( Figures 6E,F and 7C); three small hyaline patches located at costal area; R 1 single and curved apically; Rs five-branched, a terminal fork visible at right forewing of holotype ( Figure 7C), left forewing ( Figure 7B) H and 7D,E); wing with dense dark color, a large hyaline patch stretched from the middle part of wing base to middle wing, a hyaline rounded triangular spot at apex of interface between each Rs and M branches; costal area broad, Sc single, terminated at middle wing, curved apically; R 1 bifurcated into two branches at middle wing; Rs five-branched, stem Rs 1+2 more than 3 times as long as stem Rs 3+4 ; stem Rs 4a+b longer than its branches, Rs 1+2 forking at the same level as M 1+2 fork; a crossvein between R 1b and Rs 1 , basal Rs 2 and Rs 3 , Rs 3 and Rs 4a , respectively, and the crossveins inside the small spots devoid of dark color; M five-branched, M 2 two-branched, stem M 1+2 as long as stem M 3+4 and M 2a+b ; basal stem Rs 4a+b and stem M 1+2 , Rs 4b and M 1 , stem M 2a+b and M 3 connected by a crossvein, respectively; crossvein m-cua connecting M 3+4 fork and CuA; CuA and CuP straight, the crossvein between basal CuA and CuP oblique; left forewing with R 1 + Rs fork slightly proximal to M + CuA fork; right forewing with R 1 + Rs fork distinctly proximal to M + CuA fork; two straight anal veins visible.
Paratypes: NIGP200917 ( Figure 8 and Figure 19D), forewing, length 6.4 mm, width 3.0 mm, W/L 2.1; three relatively large irregular hyaline patches covered at the area of costa, subcosta and along the R 1 , a hyaline rounded triangular spot at apex of each interface between Rs 2-4 and M branches, some crossveins in spots and approximately 10 small spots at wing base; stem M 1+2 slightly longer than stem M 3+4 and twice as long as stem M 2a+b ; the three branches of M 3+4 nearly forking at one point.
Remarks. The dark color inside the forewings of the holotype is too poorly preserved to reconstruct.
Sinoagetopanorpa minuta sp. nov. (Figures 9 and 19E; ZooBank LSID urn:lsid:zoobank.org:act:512A5737-065F-49D8-BDFB-68F002A18129).  Description. Forewing with apex rounded and broad, length 4.8 mm, width 2.6 mm L/W 1.9; wing with dark color, more than 10 irregular hyaline patches focused on anterior and basal wing, some crossveins accompanied with small hyaline patches, a hyaline rounded triangular spot at apex of each interface of Rs and M1-3 branches; Sc terminated at a distance of 2/3 from wing base; R1 single and nearly straight; Rs five-branched, stem Rs1+2 length 0.6 mm, stem Rs3+4 length 0.3 mm; Rs1+2 forking at the same level as M1+2 fork Rs4 forking apically; stem Rs4a+b twice as long as its branches; a crossvein connecting basa Rs2 and Rs3; M six-branched, M2 and M4 both bifurcated into two branches, stem M1+2 as long as stem M2a+b and slightly longer than stem M3+4, stem M4a+b very short; Rs3+4 fork and stem M1+2, Rs4a+b fork and M1, stem M2a+b and M3 connected by a crossvein, respectively crossvein m-cua connecting M4a+b fork and CuA; CuA curved after the crossvein m-cua CuP straight; the crossvein between CuA and CuP robust and nearly horizontal; M + CuA fork proximal to R1 + Rs fork, R1 + Rs fork slightly proximal to the first fork of Sc; three anal veins visible, A3 short, very close to wing margin.
Etymology. The specific epithet is derived from the Latin elegans, indicating the well preserved four wings.
Diagnosis. Forewing with numerous dark-colored spots, base narrow, a hyaline rounded triangular spot at apex of each interface of Rs and M branches; Rs4a+b forked late A3 distinctly developed, the area between A3 and wing margin broad. Hind wing with Rs4a+b forked very late, R1 with three terminal branches. Etymology. The specific name is derived from the Latin minuta, small, indicating the small wing size.
Diagnosis. Forewing small, a hyaline rounded triangular spot at end of each interface of Rs and M branches; Rs 4 forking late, A 3 short, the area between A 3 and wing margin narrow.
Description. Forewing with apex rounded and broad, length 4.8 mm, width 2.6 mm, L/W 1.9; wing with dark color, more than 10 irregular hyaline patches focused on anterior and basal wing, some crossveins accompanied with small hyaline patches, a hyaline rounded triangular spot at apex of each interface of Rs and M 1-3 branches; Sc terminated at a distance of 2/3 from wing base; R 1 single and nearly straight; Rs five-branched, stem Rs 1+2 length 0.6 mm, stem Rs 3+4 length 0.3 mm; Rs 1+2 forking at the same level as M 1+2 fork; Rs 4 forking apically; stem Rs 4a+b twice as long as its branches; a crossvein connecting basal Rs 2 and Rs 3 ; M six-branched, M 2 and M 4 both bifurcated into two branches, stem M 1+2 as long as stem M 2a+b and slightly longer than stem M 3+4 , stem M 4a+b very short; Rs 3+4 fork and stem M 1+2 , Rs 4a+b fork and M 1 , stem M 2a+b and M 3 connected by a crossvein, respectively, crossvein m-cua connecting M 4a+b fork and CuA; CuA curved after the crossvein m-cua, CuP straight; the crossvein between CuA and CuP robust and nearly horizontal; M + CuA fork proximal to R 1 + Rs fork, R 1 + Rs fork slightly proximal to the first fork of Sc; three anal veins visible, A 3 short, very close to wing margin.
Etymology. The specific epithet is derived from the Latin elegans, indicating the well-preserved four wings.
Diagnosis. Forewing with numerous dark-colored spots, base narrow, a hyaline rounded triangular spot at apex of each interface of Rs and M branches; Rs 4a+b forked late, A 3 distinctly developed, the area between A 3 and wing margin broad. Hind wing with Rs 4a+b forked very late, R 1 with three terminal branches.
Material. Holotype, NIGP200921 (Figures 10 and 11), two forewings and two hind wings preserved in one specimen, interpreted as one individual. One forewing is wellpreserved, the other forewing lacking apex; two hind wings overlap with each other, poorly preserved.

Description.
Forewing with apex rounded, base obviously shrunken, the complete forewing length 5.6 mm, width 2.6 mm, L/M 2.2; dark color denser at the apical wing than basal wing, costal area with 5-6 hyaline patches; Sc terminated at a distance of 3.9 mm from wing base; R 1 single, smoothly curved near apex; pterostigma large, half below the apical R 1 ; Rs five-branched, Rs 4 bifurcated into two terminal branches, Rs 1 curved upwards, Rs 1+2 fork at the same level as M 1+2 fork, stem Rs 1+2 1.5 times as long as stem Rs 3+4 , stem Rs 4a+b long and 1.5 times as long as its branches; M six-branched, M 2 bifurcating into two branches, stem M 1+2 about 1.5-1.7 times as long as stem M 3+4 , the three branches of M 3+4 forking close, resulting in one forewing with M 3 two-branched and M 4 single, the other forewing with M 3 single and M 4 two-branched; a crossvein connecting stem Rs 3+4 and M 1+2 , stem M 2a+b and M 3 , respectively, crossvein m-cua connecting basal M 4 (or M 4b ) and CuA; CuA curved apically, CuP straight and single; M + CuA proximal to R 1 + Rs, R 1 + Rs proximal to the first fork of Sc; three anal veins, A 3 short, the area between A 3 and wing margin broad. Material. Holotype, NIGP200921 (Figures 10 and 11), two forewings and two hind wings preserved in one specimen, interpreted as one individual. One forewing is wellpreserved, the other forewing lacking apex; two hind wings overlap with each other, poorly preserved. and CuA; CuA curved apically, CuP straight and single; M + CuA + Rs proximal to the first fork of Sc; three anal veins, A3 short, t wing margin broad. Hind wing poorly preserved with dark color; costal area br as subcostal area; Sc abruptly curved to costa apically; R1 forkin apical Sc; Rs five-branched, Rs4 bifurcated into two branches, on Rs1+2 slightly longer than Rs3+4, the other hind wing with Rs1+2 nea Rs3+4; Rs4a+b fork distinctly distad to M2a+b fork; a crossvein connec M1+2; M five-branched, M2 with two branches, stem M1+2 longer th CuP single and straight, anal veins absent.
Sinoagetopanorpa grimaldii sp. nov. (Figures 12 and 19G; ZooBank LSID urn:lsid:zoobank.org:a 8253-B7D1591D0E16). Hind wing poorly preserved with dark color; costal area broad, ca. 3 times as wide as subcostal area; Sc abruptly curved to costa apically; R 1 forking near the same level as apical Sc; Rs five-branched, Rs 4 bifurcated into two branches, one hind wing with stem Rs 1+2 slightly longer than Rs 3+4 , the other hind wing with Rs 1+2 nearly twice as long as stem Rs 3+4 ; Rs 4a+b fork distinctly distad to M 2a+b fork; a crossvein connecting stem Rs 3+4 and stem M 1+2 ; M five-branched, M 2 with two branches, stem M 1+2 longer than stem M 3+4 ; CuA and CuP single and straight, anal veins absent.
Etymology. The specific name is derived from the Latin word magna, large, indic ing the large-sized wing. Etymology. The specific name is dedicated to the famous American paleoentomologist David Grimaldi.
Diagnosis. Forewing with two distinct dark stripes across the wing, with some darkcolored spots, apex of each Rs and M branches covered with a dark-colored spots and absence of the hyaline triangular spot.
Material. Four specimens, two of them with part and counterpart. Holotype, NIGP200922, a complete forewing ( Figures 12A,B and 19G). Paratypes, NIGP200923, a forewing lacking apex ( Figure 12C,D); NIGP200924, a forewing lacking wing base with part and counterpart; NIGP200925, lacking a small part of wing apex, with part and counterpart.
Description. Holotype, NIGP200922, forewing length 8.0 mm, width 4.0 mm, W/L 2.0, widest at middle wing, obviously tapering to base and apex; two dark stripes vertically lined across the wing, the larger one located at apex of R 1 , the smaller one located at Sc 3 ; many dark-colored spots scattering at wing apex, middle and basal wing, but not combined into stripe; Sc terminated at 3/4 of wing; R 1 curved apically; Rs five-branched, stem Rs 1+2 twice as long as stem Rs 3+4 , Rs 1+2 forking at the same level as M 1+2 fork, stem Rs 4a+b shorter than its branches; M six-branched, stem M 1+2 slightly longer than M 3+4 and M 2a+b ; M 3 two-branched, M 4 single, stem M 3 short; a crossvein connecting Rs 4b and M 1 , stem M 1+2 and basal Rs 4a+b , stem M 2a+b and M 3a , respectively; crossvein m-cua connecting basal M 4 and CuA; CuA single, curved apically, merged with M at the level of first Sc fork; CuP single, apically curved, the crossvein between basal CuA and CuP oblique; R 1 + Rs fork distinctly proximal to M + CuA and the first fork of Sc; two long anal veins, A 1 leaned to CuP apically, A 2 terminated at 1/3 of wing, a crossvein connecting A 1 and A 2 .
Material. Holotype NIGP200926, with part and counterpart (Figures 13 and 19H) with veins well-preserved but lacking wing apex and part of base.
Description. Forewing length 8.2 mm (as preserved), estimated length 10.0 mm width 5.0 mm, with two distinct colored stripes vertically lined across the wing, one located at apex of R1 and tapering to posterior wing, the other one located at Sc3; numerous small dark-colored spots scattering at sides of the stripes, each spot apart from the others Sc terminated at a distance of 7.5 mm from wing base; R1 curved apically; Rs fivebranched, stem Rs1+2 length 1.3 mm, stem Rs3+4 length 0. 8   Etymology. The specific name is derived from the Latin word magna, large, indicating the large-sized wing.
Diagnosis. Forewing relatively large, with two dark stripes, numerous small darkcolored spots at sides of the stripes; a distinct crossvein connecting base of Rs 1 and R 1 .
Material. Holotype NIGP200926, with part and counterpart (Figures 13 and 19H), with veins well-preserved but lacking wing apex and part of base.
Description. Forewing length 8.2 mm (as preserved), estimated length 10.0 mm, width 5.0 mm, with two distinct colored stripes vertically lined across the wing, one located at apex of R 1 and tapering to posterior wing, the other one located at Sc 3 ; numerous small dark-colored spots scattering at sides of the stripes, each spot apart from the others; Sc terminated at a distance of 7.5 mm from wing base; R 1 curved apically; Rs five-branched, stem Rs 1+2 length 1.3 mm, stem Rs 3+4 length 0.8 mm, stem Rs 4a+b length 2.2 mm, Rs 1+2 fork slightly proximal to M 1+2 fork, a crossvein connecting basal Rs 1 and R 1 ; M six-branched, stem M 1+2 length 1.4 mm, stem M 2a+b length 1.1 mm, stem M 3+4 length 1.0 mm, M 3+4 three-branched and forking at one point; stem M 1+2 and Rs 3+4 fork, stem M 2a+b and upper branch of M 3+4 connected by a crossvein, respectively; crossvein m-cua connecting lower branch of M 3+4 and CuA; CuA curved after the crossvein m-cua, CuP single and curved apically; the crossvein between basal CuA and CuP robust and nearly horizontal; two anal veins detected, A 1 leaned to CuP near apex; A 2 terminated at a distance of 4.3 mm from wing base.
Etymology. The generic name combines the Latin word rara, rare, indicating only one specimen has been found, and a mecopteran generic name Agetopanorpa. Etymology. The species name zhangi is in honor of the late paleoentomologist, Prof. Junfeng Zhang, for his contribution to Chinese paleoentomology.
Diagnosis. As for the genus. Material. Holotype, NIGP200927, a nearly complete forewing with base poorly preserved (Figures 14 and 19I).
Description. Forewing with apex somewhat rounded, length 6.4 mm, width 3.3 mm, L/W 1.9; wing with dark color, several distinct dark-colored spots at top of middle wing, costal area with four hyaline spots, a hyaline rounded triangular spot at apex of each Rs and M branches, some crossveins inside a small spot; Sc terminated distad to the level of Rs 4a+b fork; R 1 single, straight, smoothly curved apically; Rs five-branched, Rs 1 curved, stem Rs 1+2 length 1.0 mm, stem Rs 3+4 length 0.3 mm, stem Rs 4a+b as long as its branches; Type locality and horizon. Yinping Mountain, Chaohu City, Anhui Province, Chin Yinping Formation (Capitanian).
Diagnosis. Forewing moderately large in this group, Sc branches long, Sc3, Sc4, an stem Sc forming relatively small angles.
Material. Holotype, NIGP200928, a specimen lacking some part of apex, with pa and counterpart (Figures 15 and 19J). Etymology. The species name is derived from the Yinping Formation, where the specimen was collected.
Diagnosis. Forewing moderately large in this group, Sc branches long, Sc 3 , Sc 4 , and stem Sc forming relatively small angles.
Material. Holotype, NIGP200928, a specimen lacking some part of apex, with part and counterpart (Figures 15 and 19J).
Description. Forewing relatively broad, length 8.0 mm, width 4.3 mm, broadest at 2/3 of wing; wing with dense dark color, five hyaline patches at costal area, a hyaline rounded triangular spot at each apex between Rs 1 , Rs 2 and Rs 3 ; Sc terminated at a distance of 5.9 mm from wing base, the first fork of Sc at a distance of 2.3 mm from wing base; R 1 single and curved apically; Rs five-branched, stem Rs 1+2 length 1.  Material. Holotype, NIGP200929, lacking a large part of wing base, overlapped b other wing fragment at anal area, with part and counterpart (Figures 16 and 19K).
Description. Forewing covered with dense dark color, wing length 5.2 mm (as pr served), width 3.1 mm, with some small hyaline patches at costal area; a hyaline rounde triangular spot at apex of each interface between Rs and M1-3 branches; Sc3, Sc4 and Sc ste forming nearly 60-degree angles, Sc1 and Sc2 forming a 30-degree angle, each branch of S nearly the same length; R1 single and curved apically; Rs five-branched, stem Rs1+2 leng 0.7 mm, stem Rs3+4 length 0.5 mm, Rs4a+b longer than its branches, Rs1+2 forking at the sam level as M1+2 fork; a possible crossvein inside a spot connecting basal Rs2 and Rs3; M si branched, with M2 and M4 forking into two branches, stem M1+2 length 0.9 mm, stem M length 0.7 mm, stem M2a+b length 0.8 mm, stem M4a+b short; a crossvein connecting ste Rs3+4 and stem M1+2, stem M2a+b and M3; CuA curved at the apical part; other veins n preserved. Etymology. The specific name is derived from the incompletely preserved forewing. Type locality and horizon. Yinping Mountain, Chaohu City, Anhui Province, China; Yinping Formation (Capitanian).
Diagnosis. Forewing relatively small, Sc branches relatively short, Sc 3 , Sc 4 and Sc stem forming relatively large angles.
Material. Holotype, NIGP200929, lacking a large part of wing base, overlapped by other wing fragment at anal area, with part and counterpart (Figures 16 and 19K).
Description. Forewing covered with dense dark color, wing length 5.2 mm (as preserved), width 3.1 mm, with some small hyaline patches at costal area; a hyaline rounded triangular spot at apex of each interface between Rs and M 1-3 branches; Sc 3 , Sc 4 and Sc stem forming nearly 60-degree angles, Sc 1 and Sc 2 forming a 30-degree angle, each branch of Sc nearly the same length; R 1 single and curved apically; Rs five-branched, stem Rs 1+2 length 0.7 mm, stem Rs 3+4 length 0.5 mm, Rs 4a+b longer than its branches, Rs 1+2 forking at the same level as M 1+2 fork; a possible crossvein inside a spot connecting basal Rs 2 and Rs 3 ; M six-branched, with M 2 and M 4 forking into two branches, stem M 1+2 length 0.9 mm, stem M 3+4 length 0.7 mm, stem M 2a+b length 0.8 mm, stem M 4a+b short; a crossvein connecting stem Rs 3+4 and stem M 1+2 , stem M 2a+b and M 3 ; CuA curved at the apical part; other veins not preserved.
Hind wings of Sinoagetopanorpidae fam. nov.
The hind wings of Sinoageotpanorpidae fam. nov. are often preserved as isolated wings. Fourteen isolated hind wings (NIGP200930, NIGP200931, NIGP200932, NIGP200933, NIGP200934, NIGP200935, NIGP200936, NIGP200937, NIGP200938, NIGP200939, NIGP200940, NIGP200941, NIGP200942 and NIGP200943) were found (four with part and counterpart; NIGP200932 preserved a pair of hind wings). Two hind wings accompanied with forewings (S. lini. sp. nov. and S. elegans sp. nov.), and the rest of hind wings remain elusive in terms of their systematic positions. Similar to forewings, the hind wings are very stable in venation, with two types of venational patterns distinguished. The most common one (13/14) with three-branched R 1 is represented by S. elegans sp. nov., but the hind wings of S. elegans sp. nov. are too incomplete with poor preservation to compare with other isolated hind wings; thus, no isolated hind wings are reluctantly attributed to S. elegans sp. nov. The rare one (one specimen) with two-branched R 1 is represented by S. lini sp. nov.
The specimen NIGP200930 ( Figures 17A,B and 18A) is a well-preserved hind wing and described as follows: wing broad, base narrow, broadest at near middle wing, length 6.6 mm, width 3.6 mm, L/W 1.8; wing apex with dark color, costal area with three hyaline patches, basal one oval shaped and large; basal wing devoid of dark color; lines of dark color along longitudinal veins; costal area broad, ca. 3 times as wide as subcostal area; Sc abruptly curved to costa near apex; R 1 with three evenly developed terminal branches, R 1b paralleled to R 1c , forking near the level of apical Sc; Rs five-branched, with Rs 4 bifurcated into two branches, stem Rs 1+2 more than twice as long as stem Rs 3+4

Discussion
Sinoagetopanorpidae fam. nov. resemble members of the subfamily Agetopanorpinae of Permochoristidae in venation. Both groups are characterized by the following venational characteristics: Sc usually with three evenly developed elongated branches, Rs with five branches and M generally with six branches. However, it is very conspicuous that sinoagetopanorpids possess broad oval-shaped forewings with broad costal area, and the forking patterns of the three branches of the M 3+4 forks are varied: M 4 bifurcating into two branches and M 3 single or M 3 bifurcating into two branches and M 4 single, or even three branches of M 3+4 forking at the same point. With the consideration of these different forewing characteristics, we erected the new family Sinoagetopanorpidae fam. nov.
Sinoagetopanorpidae resemble Choristopsychidae Martynov, 1937 in having broad oval-shaped wings: Sinoagetopanorpidae possess a wing aspect ratio of 1.7-2.2, while 1.5-2.0 in Choristopsychidae, and both families show a broad costal area with three evenly developed branched Sc. Choristopsychidae was placed by many authors in the Permochoristidae family for its venational similarity with Agetopanorpinae [45][46][47]; however, Qiao et al. [48] discovered numerous exquisite specimens from the Middle−Upper Jurassic (originally assigned to the Middle Jurassic) Daohugou biota, and erected the family Choristopsychidae. Sinoagetopanorpidae differ from Choristopsychidae in a combination of the following characteristics: in terms of forewings, Sinoagetopanorpidae have at least fivebranched Rs instead of four-branched, six-branched M with three-branched M 3+4 instead of five-branched M with two-branched M 3+4 ; in addition, the hind wings of Sinoagetopanorpidae have single Sc instead of two-branched Sc; furthermore, Sinoagetopanorpidae are confined to the Permian strata, whereas Choristopsychidae occurred in the Jurassic.
Based on our study of abundant new specimens, we found that the venational pattern of Sinoagetopanorpidae is rather stable: Sc generally with three branches (except the two new species of Permoagetopanorpa); Rs five-branched; M with six branches where M 2 and M 4 (or M 3 ) bifurcate into two branches (except for Raragetopanorpa zhangi sp. nov., which possesses a five-branched M with two-branched M 2 and a single M 4 ) and anal veins generally with two branches, but A 3 detected in S. elegans and S. minuta. To distinguish these species, dark-colored pattens play another important role in species-level classification. In the forewings, four kinds of colored patterns are recognized: (1) the absence of a hyaline rounded triangular spot at the apex of each interface between Rs and M branches; (2) the apex of each Rs and M vein has a dark-colored spot and two dark-colored stripes across the wing; (3) a hyaline rounded triangular spot at the apex of each interface between Rs and M branches, the wing with a dense dark color; and (4) a hyaline rounded triangular spot at the apex of each interface between Rs and M branches, the wing with numerous colored spots.
Our discovery of two specimens preserved with both forewings has some implications for venational variations. The holotype of S. lini sp. nov. possesses the right forewing with Rs 1 armed with a terminal fork, while the left forewing with a single Rs 1 . The variation of the terminal fork in the anterior branches of Rs in one individual can be found in other mecopteran families, such as Cimbrophlibiidae [49] and Panorpidae [50,51]. The two forewings of holotype of S. elegans sp. nov. show variation in the forking pattern of the three branches of M 3+4 : one forewing with M 3 two-branched and M 4 single, but the other with M 4 two-branched and M 3 single, indicating that the forking pattern of M 3+4 is unstable. Therefore, we do not regard these characters as an interspecific diagnostic characteristic.
During the Capitanian, the fossil locality was near 32 • N on the Northeast Yangtze Platform in the eastern Paleotethys, which split from the Gondwana supercontinent during the Silurian [44,52], and it possibly had a lagoonal paleoenvironment under the large-scale regression [35,38]. The extant known scorpionflies, with no exception, are weak flyers with low dispersal capacity. Therefore, the endemic mecopteran group Sinoagetopanorpidae fam. nov., the representative mecopterans in the Yinping Formation, might have evolved independently on the Yangtze Platform. The new family from the late Capitanian might have become extinct during the end-Guadalupian mass extinction, which was possibly associated with large-scale volcanic activities [53,54].