Mesoraphidia daohugouensis, Lü, Ya-Nan, Liu, Xingyue & Dong, Ren, 2015

Lü, Ya-Nan, Liu, Xingyue & Dong, Ren, 2015, First record of the fossil snakefly genus Mesoraphidia (Insecta: Raphidioptera: Mesoraphidiidae) from the Middle Jurassic of China, with description of a new species, Zootaxa 3999 (4), pp. 560-570 : 561-568

publication ID	https://doi.org/ 10.11646/zootaxa.3999.4.6
publication LSID	lsid:zoobank.org:pub:B239D382-BC87-4D4F-920D-24496F4D2080
DOI	https://doi.org/10.5281/zenodo.6097365
persistent identifier	https://treatment.plazi.org/id/03961C5E-FFF6-5419-609D-E467FD98FACF
treatment provided by	Plazi
scientific name	Mesoraphidia daohugouensis
status	sp. nov.

Treatment

Mesoraphidia daohugouensis View in CoL sp. nov.

( Figs. 1–11 View FIGURES 1 – 3 View FIGURES 4 – 5 View FIGURES 6 – 8 View FIGURES 9 – 10 View FIGURE 11 )

Diagnosis. The new species can be distinguished by the following combination of features. Head ovoid. Antennae long, flagellomere nearly as long as wide. Pronotum slightly shorter than head. Forewing narrowly elliptical. Single proximal sc-r crossvein. Pterostigma elongate, gradually darkened distad, respectively closed by a costal crossvein proximally and veinlet of R distally. Pterostigmal crossvein absent. Two nearly equal long, elongate radial cells. Rs with two branches. MA with two simple branches. Single, elongate discoidal cell (doi) in hindwing.

Description of male holotype. Body length 7.5 mm. Head ovoid, 1.4 mm long, 1.2 mm wide. Antennae with 31 flagellomeres visible (more than 36 flagellomeres by estimation); scape inflated and pedicel not, flagellomere nearly as long as wide. Pronotum subquadrate, 1.2 mm long, 1.0 wide, slightly shorter than head. Only right foreleg incompletely preserved with tarsomeres fragmentarily visible.

Forewing narrowly elliptical, 6.7 mm long, 2.2 mm wide. Costal area slightly dilated at proximal 1/6, with seven crossveins. Sc running into C before base of pterostigma. Subcostal area nearly half of maximum width of costal area. Single, proximal sc-r present. Pigmented pterostigma well developed, elongate, 2.3 mm long, terminating at veinlet of R, gradually darkened distad, proximally closed by a costal crossvein. R slightly curved distally, with only one simple veinlet, which is distinctly inclined toward wing apex. Rs+MA originating from R nearly at proximal 1/3. Rs with one marginally forked anterior branch and a simple posterior branch. MA distally forked. Two radial cells (1 r and 2 r) present between R and Rs; 2 r slightly shorter than 1 r, but nearly twice as wide as 1 r. Single discal cell (dc) present. Two crossveins present between MA and MP, forming two long medial cells (1 m and 2 m). MP diverging from CuA with a short distance to diverging point of M+CuA+R, with two main branches, anterior branch distally trifurcated, posterior branch distally bifurcated. Three discoidal cells (doi) posterior to MP in the forewing (forming a triangle). CuA with three simple branches. CuP simple. Two cua-cup crossveins present.

Hindwing 5.9 mm long, 1.7 mm wide. Costal area nearly half of width of forewing costal area, with seven crossveins. Sc fused into C slightly beyond mid-point of wing. R slightly curved posteriad distally, with one short distal veinlet, which is distinctly inclined toward wing apex. Pterostigma well developed, elongate (ca. 2.2 mm long), gradually darkened distad, proximally closed by a costal crossvein. Rs with one marginally forked anterior branch and a simple posterior branch. MA distally forked. Three crossveins present between R and Rs, forming three radial cells respectively; 3 r conspicuously longer than other radial cells, nearly 2 times as long as 1 r, about 2.5 times as long as 2 r. Single, long dc present between Rs and MA. MA originating from R posterior to branching point of MP from R and coalescent with Rs for distance nearly 1/5 length of forewing. MP deeply forked, both anterior branch and posterior branch distally trifurcated.

Abdomen 3.7 mm long, 1.1 mm wide at maximum in dorsal view; color of abdomen paler than metathorax; segments dark and joints between segments pale; 4–6 segments longer than others; abdominal apex much narrower than pregenital segments; a valvate sclerite (gonocoxite 9?) preserved.

Description of female paratypes. Body 8.8–11.1 mm long (exclude ovipositor). Ovipositor long, 2.0– 3.6 mm long, 0.3 mm wide. Head ovoid. Pronotum about 1.0 mm long, slightly shorter than head. Meso- and metathorax poorly preserved; details not visible. One fore leg and two hind legs incompletely preserved covered by dense hair, fore femur 0.6 mm, tibia 1.4 mm, hind femur 1.4 mm, tibia 1.8 mm, tarsal claw of hind leg visible. Forewing ovoid, 8.3–8.7 mm long, nearly 3.0 wide. Hindwing 7.0– 7.9 mm long, 2.4–2.7 mm wide. Fore- and hindwing slightly broader than those in male. Venation almost identical to that in holotype. In paratype (CNU-RAP-NN-2015003) an ovoid anal cell (ac) preserved.

Derivation of name. The new species is named after its type locality, Daohugou, in Inner Mongolia, China.

TABLE 1. List of known species of Mesoraphidiidae

Age Subamily/tribe Species Locality Reference

Upper Cretaceous Alloraphidiinae Alloraphidia kyzylzharica Makarkin & Khramov, 2014 Kazakhstan Makarkin & Khramov 2014

Upper Cretaceous Nanoraphidiini Grimaldiraphidia luzzii (Grimaldi, 2000) New Jersey amber, USA Bechy & Wolf-Schwenninger 2011

Upper Cretaceous Nanoraphidiini Nanoraphidia electroburmica Engel, 2002 Burmese amber, Engel 2002 Myanmar

Middle Cretaceous Alloraphidiinae Alloraphidia dorfi Carpenter, 1967 Canada Carpenter 1967

Lower Cretaceous Alloraphidiinae Alloraphidia anomala Ren, 1997 China Ren 1997

Lower Cretaceous Alloraphidiinae Alloraphidia petrosa Ponomarenko, 1988 Mongolia Ponomarenko 1988

Lower Cretaceous Alloraphidiinae Archeraphidia hosbayari Ponomarenko, 1988 Mongolia Ponomarenko 1988

Lower Cretaceous Alloraphidiinae Archeraphidia yakowlewi Ponomarenko, 1988 Mongolia Ponomarenko 1988

Lower Cretaceous Alloraphidiinae Pararaphidia deserta (Ponomarenko, 1988) Mongolia Willmann 1994

Lower Cretaceous Mesoraphidiinae Alavaraphidia imperterrita Pérez-de la Fuente, Peñalver, Delclòs & Spanish amber, Spain Pérez-de la Fuente et al. 2012

Engel, 2012

Lower Cretaceous Mesoraphidiinae Amarantoraphidia ventolina Pérez-de la Fuente, Peñalver, Delclòs & Spanish amber, Spain Pérez-de la Fuente et al. 2012

Engel, 2012

Lower Cretaceous Mesoraphidiinae Cretinocellia cellulosa Ponomarenko, 1988 Mongolia Pérez-de la Fuente et al. 2012

Lower Cretaceous Mesoraphidiinae Huaxiaraphidia shandongensis Hong, 1992 China Hong 1992a

Lower Cretaceous Mesoraphidiinae Huaxiaraphidia sinensis Hong, 1992 China Hong 1992a

Lower Cretaceous Mesoraphidiinae Jilinoraphidia dalaziensis Hong & Chang, 1989 China Hong & Chang 1989

Lower Cretaceous Mesoraphidiinae Kezuoraphidia kezuoensis (Hong, 1992) China Willmann 1994

Lower Cretaceous Mesoraphidiinae Mesoraphidia amoena Ren, 1997 China Ren 1997

Lower Cretaceous Mesoraphidiinae Mesoraphidia durlstonensis Jepson, Coram & Jarzembowski, 2009 England Pérez-de la Fuente et al. 2012

Lower Cretaceous Mesoraphidiinae Mesoraphidia ednae Jepson, Coram & Jarzembowski, 2011 England Jepson et al. 2011

Lower Cretaceous Mesoraphidiinae Mesoraphidia furcivenata Ren in Ren et al., 1995 China Ren et al. 1995

Lower Cretaceous Mesoraphidiinae Mesoraphidia glossophylla ( Ren, 1997) China Engel 2002

Lower Cretaceous Mesoraphidiinae Mesoraphidia heteroneura Ren, 1997 China Pérez-de la Fuente et al. 2012

Lower Cretaceous Mesoraphidiinae Mesoraphidia hilli Jepson, Coram & Jarzembowski, 2011 England Jepson et al. 2011

……continued on the next page TABLE 1. (Continued)

Age Subamily/tribe Species Locality Reference

Lower Cretaceous Mesoraphidiinae Mesoraphidia koreenensis Engel, Lim & Baek, 2006 Korea Engel et al. 2006

Lower Cretaceous Mesoraphidiinae Mesoraphidia longistigmosa ( Ren, 1994) China Engel 2002

Lower Cretaceous Mesoraphidiinae Mesoraphidia myrioneura ( Ren, 1997) China Engel 2002

Lower Cretaceous Mesoraphidiinae Mesoraphidia obliquivenatica ( Ren, 1994) China Engel 2002

Lower Cretaceous Mesoraphidiinae Mesoraphidia phantosma Engel, Lim & Baek, 2006 Korea Engel et al. 2006

Lower Cretaceous Mesoraphidiinae Mesoraphidia polyphlebia ( Ren, 1994) China Engel 2002

Lower Cretaceous Mesoraphidiinae Mesoraphidia shangyuanensis ( Ren, 1994) China Engel 2002

Lower Cretaceous Mesoraphidiinae Mesoraphidia sinica Ren, 1997 China Ren 1997

Lower Cretaceous Mesoraphidiinae Mesoraphidia websteri Jepson, Coram & Jarzembowski, 2009 England Jepson et al. 2009

Lower Cretaceous Mesoraphidiinae Necroraphidia arcuata Pérez-de la Fuente, Peñalver, Delclòs & Engel, Spanish amber, Spain Pérez-de la Fuente et al. 2012

2012

Lower Cretaceous Mesoraphidiinae Proraphidia gomezi Jepson & Jarzembowski, 2008 Spain Jepson & Jarzembowski

Lower Cretaceous Mesoraphidiinae Proraphidia hopkinsi Jepson & Jarzembowski, 2008 England Jepson & Jarzembowski 2008

Lower Cretaceous Mesoraphidiinae Siboptera fornicata ( Ren, 1994) China Ren 1997

Lower Cretaceous Mesoraphidiinae Xuraphidia liaoxiensis Hong, 1992 China Hong 1992b

Lower Cretaceous Mesoraphidiinae Yanoraphidia gaoi Ren , in Ren et al. 1995 China Pérez-de la Fuente et al. 2012

Lower Cretaceous Nanoraphidiini Cantabroraphidia marcanoi Pérez-de la Fuente, Nel, Peñalver & Spanish amber, Spain Pérez-de la Fuente et al. 2010

Delclòs, 2010

Lower Cretaceous Nanoraphidiini Grimaldiraphidia mitchelli ( Jepson, Coram & Jarzembowski, 2009) England Bechy & Wolf-Schwenninger 2011

Lower Cretaceous Nanoraphidiini Grimaldiraphidia purbeckensis (Jepson, Coram & Jarzembowski, England Bechy & Wolf-Schwenninger 2011

2009)

Lower Cretaceous Nanoraphidiini Lebanoraphidia nana Bechly & Wolf-Schwenninger, 2011 Lebanon Bechy & Wolf-Schwenninger 2011

Lower Cretaceous Nanoraphidiini Nanoraphidia lithographica Jepson, Coram & Jarzembowski, 2011 Spain Jepson et al. 2011

Lower Cretaceous Ororaphidiinae Styporaphidia ? hispanica Pérez-de la Fuente, Peñalver, Delclòs & Spanish amber, Spain Pérez-de la Fuente et al. 2012

Engel, 2012

Lower Cretaceous Mesoraphidiinae Iberoraphidia dividua Jepson, Coram & Jarzembowski, 2011 Spain Jepson et al. 2011

……continued on the next page TABLE 1. (Continued)

Subamily/tribe Species Locality Reference Upper Jurassic- Lower Alloraphidiinae Alloraphidia asiatica Ponomarenko, 1993 Russia Ponomarenko 1993 Cretaceous

Upper Jurassic- Lower Alloraphidiinae Archeraphidia baisensis Ponomarenko, 1993 Russia Ponomarenko 1993 Cretaceous

Upper Jurassic- Lower Alloraphidiinae Pararaphidia vitimica ( Martynova, 1961) Russia Engel 2002 Cretaceous

Upper Jurassic- Lower Mesoraphidiinae Baisoraphidia glossopteron Ponomarenko, 1993 Russia Ponomarenko 1993 Cretaceous

Upper Jurassic- Lower Mesoraphidiinae Baisoraphidia orientalis Ponomarenko, 1993 Russia Ponomarenko 1993 Cretaceous

Upper Jurassic- Lower Mesoraphidiinae Mesoraphidia medialis (Ponomarenko, 1993) Russia Bechy et al. 2011 Cretaceous

Upper Jurassic- Lower Mesoraphidiinae Siboptera eurydictyon Ponomarenko, 1993 Russia Ponomarenko 1993 Cretaceous

Upper Jurassic- Lower Mesoraphidiinae Sinoraphidia viridis Hong, 1982 China Hong 1982 Cretaceous

Upper Jurassic Mesoraphidiinae Mesoraphidia elongata Martynov, 1925 Kazakhstan Martynov 1925 Upper Jurassic Mesoraphidiinae Mesoraphidia gobiensis Ponomarenko, 1988 Mongolia Ponomarenko 1988 Upper Jurassic Mesoraphidiinae Mesoraphidia grandis Martynov, 1925 Kazakhstan Martynov 1925 Upper Jurassic Mesoraphidiinae Mesoraphidia inaequalis Martynov, 1925 Kazakhstan Martynov 1925 Upper Jurassic Mesoraphidiinae Mesoraphidia pterostigmalis Martynova, 1947 Kazakhstan Martynov 1947 Upper Jurassic Mesoraphidiinae Mesoraphidia similis Martynov, 1925 Kazakhstan Martynov 1925 Upper Jurassic Mesoraphidiinae Proraphidia turkestanica Martynova, 1947 Kazakhstan Martynov 1947 Upper Jurassic Nanoraphidiini Grimaldiraphidia parvula ( Martynov, 1925) Kazakhstan Bechy & Wolf-Schwenninger 2011 Middle Jurassic Mesoraphidiinae Mesoraphidia daohugouensis sp. nov. China Middle Jurassic Ororaphidiinae Ororaphidia megalocephala Engel & Ren, 2008 China Engel & Ren 2008 Middle Jurassic Ororaphidiinae Styporaphidia magia Engel & Ren, 2008 China Engel & Ren 2008 Type material. Holotype, CNU-RAP-NN-2015001, male, a well preserved specimen in dorsal aspect, with left fore- and hindwings largely not preserved. Paratype, CNU-RAP-NN-2015002p/c, female, a well preserved complete specimen in dorsal aspect. Paratype, CNU-RAP-NN-2015003; female, a nearly complete specimen in lateral view.

Type locality and horizon. All collected from Daohugou locality (41°18’30”N, 119°13’00”E), Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Middle Jurassic (Aalenian-Bajocian).

Remarks. The placement of this new species in the genus Mesoraphidia is based on the configuration of the following characters: (1) size, forewing length 6.7–8.7 mm [species with forewing length shorter than 6 mm are transformed to or put in the tribe Nanoraphidiini and the genus Amarantoraphidia]; (2) antenna with flagellomere about as long as wide [flagellomere 1.5 times as long as wide in Alavaraphidia]; (3) presence of single proximal sc-r crossvein [none in Cretinocellia ; two sc-r crossveins in Xuraphidia ; numerous in Sinoraphidia (this feature may be artificial)]; (4) pterostigma elongate [pterostigma shorter in Sinoraphidia ; strongly shortened in Proraphidia ], gradually darkened distad, respectively closed by a costal crossvein proximally and veinlet of R distally [pterostigma with a diffuse base in the subfamily Ororaphidiinae and the genus Necroraphidia; pterostigma extending distally to apex of R in Yanoraphidia ]; (5) only one R veinlet [three in Cretinocellia ]; (6) Rs+MA originating from R at about proximal 1/3 [Rs+MA originating from R at proximal 1/ 2 in Siboptera ]; (7) MP diverging from CuA with a short distance to diverging point of M+CuA+R [apparent triadic branching of R, M, and CuA in subfamily Alloraphidiinae]; (8) presence of two radial cells [three radial cells present in Baissoptera , Jilinoraphidia , Kezuoraphidia and Xuraphidia ]; (9) no crossvein between MP posterior main branch and CuA (several crossveins there and in vannal region in Huaxiaraphidia ).

The new species differs from other described species of Mesoraphidia based on the following characters: (1) medium body-size, forewing length 6.7–8.7 mm [e.g. forewing of M. glossophylla , M. grandis , M. inaequalis , M. longistigmosa , M. myrioneura , M. obliquivenatica , M. polyphlebia and M. similis , approximately two times longer than the new species, or even longer]; (2) pronotum slightly shorter than head [e.g. pronotum longer than head in M. shangyuanensis , while strongly shortened, nearly half of head length in M. gobiensis ]; (3) single veinlet of R [e.g. three in M. inaequalis and M. similis ]; (4) pterostigma closed proximally by a costal crossvein [e.g. a diffuse base in M. amoena and M. durlstonenesis ]; (5) pterostigma closed distally by one veinlet of R [e.g. a row of oblique transverse crossveins in distal portion of pterostigma in M. elongata ]; (6) absence of pterostigmal crossvein [e.g. one pterostigmal crossvein present in M. ednae , M. glossophylla , M. longistigmosa and M. medialis ]; (7) presence of two forewing radial cells [e.g. 3–4 forewing radial cells present in M. koreenensis M. myrioneura , M. phantosma and M. sinica ,]; (8) the length of 1 r nearly equal to 2 r [e.g. 1 r about 1.5 times as long as 2 r in M. heteroneura ]; (9) Rs with less than three branches [e.g. Rs with 4–5 branches in M. grandis , M. hilli , M. obliquivenatica , M. polyphlebia and M. pterostigmalis ]; (10) hindwing Sc fused into C at about mid-point of wing [e.g. Sc terminates at two-thirds of wing length from wing apex in M. websteri ]; (11) presence of single medial cell in hindwing [e.g. two medial cells present in M. furcivenata ].