Liberitoma compta Sun, Rasnitsyn, Zhuang & Shih, 2024

Sun, Chenhui, Rasnitsyn, Alexandr P., Wedmann, Sonja, Zhuang, Jialiang, Shih, Chungkun, Ren, Dong & Gao, 2024, Revisions * new taxa * and venation transformations of the sawfly family Blasticotomidae sensu lato (Hymenoptera: ºenthredinoidea) highlight the evolution of the basal Hymenoptera, Zoological Journal of the Linnean Society 202 (1) : -

Liberitoma compta Sun, Rasnitsyn, Zhuang & Shih sp. nov.

( Figs 4, 5)

http://zoobank.org/urn:lsid:zoobank.org:act:6E1D3226-

993D-4E53-9672-53DEB11C4A8C

Diagnosis: Antenna with thread at least one-third of first flagellomere* with five preserved segments* each with length to width ratio ~2:1. Forewing with Sc basal branch short* vertical* meeting R prior to 1-M base; Sc 2 short* slightly proclined* merging into R at level of 1-M base; Sc 1 proclined and short (about as long as Sc 2).

Etymology: The species name is derived from the Latin word ‘comptus’* meaning ornamented* adorned* referring to this sawfly being preserved in good condition.

Material: Holotype * CNU-HYM-NN2022302. Paratypes: CNU-HYM-NN2022303; CNU-HYM-NN2022304; CNU-HYM-NN2022305; CNU-HYM-NN2022306.

Locality and horizon: Latest Middle Jurassic (late Callovian)* Jiulongshan Formation ; Daohugou Village * Wuhua ºownship* Ningcheng County * Chifeng City * Inner Mongolia * China. Description: Holotype * CNU-HYM-NN2022302 ( Fig. 4). Male. Body length 13.4 mm as preserved (excluding antennae). Surface coarsely punctated* especially on head* thorax* and the first abdominal segments ( Fig. 4A * B).

Head: Short* ovate* with eyes not small* roundish* apparently bulging* placed near the fore margin of head. Mandibles with two teeth visible.

Antenna: First flagellomere (2.1 mm) shorter than width of head (2.8 mm)* and about three times as long as thread. Thread with five segments preserved* thicker than those of type species* length (in millimetres) of each flagellomere: 0.16* 0.15* 0.15* and 0.13 (the last flagellomere was not measured owing to partial preservation)* length to width ratio ~2:1 ( Fig. 4C).

Forewing: Sc with basal branch short* vertical; Sc 2 short* slightly proclining; Sc 1 short* oblique* basal branch meeting R well basal to 1-M base; Sc 2 merging into R basal to 1-Rs for more than 1-Rs length; Sc 1 curved* about as long as Sc 2 ( Fig. 4D). Pterostigma fully sclerotized. Vein M+Cu slightly bent; 1-M (0.9 mm) obviously curved* about three times as long as 1-Rs (0.3 mm). Crossvein 2r-rs (0.8 mm) roughly two times as long as 1r-rs (0.3 mm)* oblique and intersecting with pterostigma near its apex; 2r-m connecting with cell 2r almost at the end of the cell* and distinctly bent towards apex of wing at middle. Cell 1r large; 2r and 2rm narrow and equal in lengths and widths.

Hind wing: Sc two-branched* forking far from apex of costal space* with hind branch very short and fore one longer* oblique ( Fig. 4E). Basal hamuli visible in left hind wing.

Legs: Long and narrow* fore tibia 3.0 mm long* tarsus relatively shorter than that in type species* tarsus with two segments present; tarsomere I (1.42 mm) longer than tarsomere II (0.83 mm)* claw strongly bent. Middle tibia 3.4 mm long* with two apical spurs visible.

Abdomen: Male genitalia apparently not reverted* with gonocoxa and gonostylus massive* gonostylus apparently with gonomacula* volsella with stick-like digitus* cuspis and penis valves not distinct.

Remarks: Gonomacula has not been recorded in other ºenthredinoidea yet including extant Blasticotomidae .

Paratype CNU-HYM-NN2022303 ( Fig.5A, B)

Sex unknown. Generally similar to the holotype as preserved. Antenna well preserved and complete. First flagellomere (1.9 mm) obviously shorter than width of head (2.6 mm)* approximately two times as long as thread* and a total of seven thread segments preserved; length to width ratio of each flagellomere about two ( Fig. 5A). Only left forewing partly preserved. Vein Sc three-branched* the first branch uniquely running into R very close to separation of 1-M from M+Cu; Sc 1 preserved incompletely; Sc 2 oblique* meeting R before origin of 1-Rs ( Fig. 5B) .

Paratype CNU-HYM-NN2022304 ( Fig. 5C–E)

Sex unknown. Similar to the holotype. Mandibles incompletely preserved* thick and curved ( Fig. 5D). Legs partly well preserved* all with vaguely visible numerous fine spines. Right foreleg: tibia (2.2 mm) with two apical spurs present* almost equal in length; tarsus with four segments present* tarsomere I longer than tarsomeres II and III combined; lengths of individual tarsomeres (in millimetres): 1.01* 0.43* 0.36* and 0.34. Right middle tibia length 2.8 mm * tarsomeres I and V longer than tarsomeres II* III* and IV combined; lengths of individual tarsomeres (in millimetres): 1.24* 0.49* 0.42* 0.27* and 0.55. Right hind tibia 4.4 mm long* with no spur preserved. Left hind tibia (4.2 mm) with two apical spurs preserved* one long and one short; tarsus with five tarsomeres and one tarsal claw preserved* tarsomere I approximately equal to tarsomeres II* III* and IV combined; lengths of individual tarsomeres (in millimetres): 1.24* 0.51* 0.40* 0.35* and 0.67. Body length 9.9 mm * forewing length 12.7 mm .

Paratype CNU-HYM-NN2022305 ( Fig. 5F, G)

Female. Forewings both partly preserved. Vein Sc triple-branched* shaped like an inverted ‘Y’; the first branch intersecting with R before separation of 1-M from M+Cu; Sc 1 and Sc 2 both oblique in different directions; Sc 1 (0.28 mm) ~1.7 times as long as Sc 2 (0.16 mm)* merging into C a short distance before separation of 1-Rs from R; Sc 2 meeting R before origin of 1-Rs ( Fig. 5G). M+Cu slightly bent; 1-M (1.1 mm) curved and about three times more than 1-Rs (0.3 mm). Crossvein 1r-rs obviously shorter than 1r-m; 2r-rs (1.3 mm) roughly three times as long as 1r-rs (0.4 mm)* oblique and intersecting with pterostigma near its apex. Cell 1r large; cell 2r and cell 2rm narrow. Body length 16.9 mm * right forewing length 13.3 mm * ovipositor length 3.0 mm.

Paratype CNU-HYM-NN2022306 ( Fig.5H–J)

Male. Body length 16.5 mm (excluding antennae). Mandible paired preserved* bent like a sickle ( Fig. 5I). Forewings partly preserved* Vein Sc with three branches ( Fig. 5J)* the first branch merging into R before separation of 1-M from M+ Cu; Sc 1 (0.31 mm) short* ~1.9 times as long as Sc 2 (0.16 mm)* and inclining towards apex of the wing* merging into C before separation of 1- Rs from R; Sc 2 slightly oblique* meeting R before origin of 1- Rs. M + Cu slightly bent* 1-M (1.1 mm) arcuately* and approximately three times more than 1- Rs (0.3 mm). Cell 1r large; 1mcu long and lengthening to posterior margin. Male genitalia different from holotype * with gonocoxa longer* gonostylus shorter and much narrower. Body length 16.7 mm * right forewing length 10.3 mm * genitalia length 2.2 mm .

Liberitoma incompleta Sun, Rasnitsyn, Zhuang & Shih sp. nov.

( Figs 6, 7)

http://zoobank.org/urn:lsid:zoobank.org:act:ACA2BD6F-

3B75-4160-9321-ADE470B52CCE

Diagnosis: Forewing with basal branch of Sc vertical* meeting R distinctly before level of 1-M base; Sc 2 obviously proclined and short* meeting R distinctly behind level of 1-M base; and Sc 1 curved and long.

Etymology: The species name is derived from the Latin word ‘incompletus’* meaning incomplete and referring to fragmentary preservation of this fossil sawfly.

Material: Holotype * CNU-HYM-NN2022307. Paratypes: CNU-HYM-NN2022308; CNU-HYM-NN2022309.

Locality and horizon: Latest Middle Jurassic (late Callovian)* Jiulongshan Formation; Daohugou Village* Wuhua ºownship* Ningcheng County* Chifeng City* Inner Mongolia * China.

Description: Holotype * CNU-HYM-NN2022307 ( Fig. 6). Female. Body length 14.3 mm as preserved (excluding antennae). Head* thorax* and the first abdominal segment coarsely punctated.

Head: Eye big* flat* occupying most of head side* distant from fore margin of head. Antenna not preserved.

Forewing: Only left forewing partly preserved. Vein Sc with basal branch vertical* short; Sc 2 proclined* short; Sc 1 curved* long; basal branch of Sc meeting R slightly before level of 1-M base; Sc 2 merging into R at midway between basal branch of Sc and 1-Rs; Sc 1 almost reaching level of 1-Rs ( Fig. 6C). Pterostigma fully sclerotized. M+Cu slightly bent* 1-M (1.1 mm) bent* about five times as long as 1-Rs (0.2 mm) ( Fig. 6C). 1-Cu bent* with short adventitious veinlet within cell 1cu ( Fig. 6D). Cell 1mcu distinctly widened apically* oblique and long. Crossvein 1r-rs obviously shorter than 2-RS; 2r-rs (0.9 mm) about two times as long as 1r-rs (0.4 mm)* oblique and meeting with pterostigma near its apex; 2r-m connecting with 2r roughly at three-quarters of its length. Cells 1r large* 2r and 2rm narrow and nearly equal in width* 1mcu long* and 2mcu wide.

Hind wing: Sc with two branches* forking far from apex of costal space* with hind branch very short and fore one longer* oblique.

Legs: Poorly preserved* except for hind femora and tibiae 4.1 mm and 3.7 mm long* respectively.

Abdomen: Ovipositor short ( Fig. 6E)* 2.3 mm long; saw sheaths wide* subtriangular* slightly longer than wide.

Paratype, CNU-HYM-NN2022308 ( Fig.7A, B)

Female: Body length 12.6 mm as preserved ( Fig. 7A). Vein Sc three-branched in forewing* the first branch of Sc running into R much prior to separation of 1-M from M+Cu; Sc 1 (0.43 mm) relatively extended compared with Sc 2 (0.17 mm)* ~2.5 times as long as Sc 2 * and inclining to apex of the wing* merging into C a short distance before separation of 1-Rs from R * shorter than holotype; Sc 2 vertical* sub-parallel to the first branch* meeting R before origin of 1-Rs ( Fig. 7B). M+Cu slightly bent; 1-M (1.0 mm) with curvature about three times more than 1-Rs (0.3 mm). Crossvein 1r-rs obviously shorter than 1r-m; 2r-rs (1.0 mm) about two times more than 1r-rs (0.4 mm)* oblique and meeting with pterostigma nearly its apex; 2r-m connecting with cell 2r at three-quarters of its length* and slightly bent towards apex of wing at middle. Cell 1r large* 2r narrow. Body length 12.6 mm * forewing length 11.0 mm* ovipositor length 1.7 mm .

Paratype, CNU-HYM-NN2022309 ( Fig.7C, D)

Sex unknown: Body length 10.4 mm * with head not preserved and abdomen partly preserved ( Fig. 7C). Forewings partly preserved. Vein Sc with three branches* the first branch intersecting with R a short distance before separation of 1-M from M+Cu; Sc 1 (0.33 mm) relatively extended compared with Sc 2 (0.12 mm)* ~2.7 times as long as Sc 2 and bent to apex of the wing* merging into C before separation of 1-Rs from R; Sc 2 marginally oblique and meeting R before origin of 1-Rs ( Fig. 7D). M+Cu slightly bent; 1-M (1.1 mm) curved* and about four times as long as 1-Rs (0.26 mm). Crossvein 2r-rs (0.9 mm) about two times as long as 1r-rs (0.4 mm)* oblique and intersecting with pterostigma near its apex; 2r-m adjoining with cell 2r at three-quarters of its length* and bent towards apex of wing at middle. Cell 1r large* 2r and 2rm narrow and equal in width. Body length 10.4 mm * right forewing length 10.5 mm.

Aduantoma Sun, Rasnitsyn, Zhuang & Shih gen. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:F2C9AEB7-

D2BA-4CC8-A48A-E6724C7657D7

Type species: Aduantoma insolita Sun * Rasnitsyn* Zhuang & Shih sp. nov.

Etymology: The generic name is a combination of the Latin ‘Aduant-’* meaning associated* and referring to the Sc in forewing* and ‘-toma’ from the type genus Undatoma Rasnitsyn * 1977. Gender feminine.

Included species: ºype species only.

Diagnosis: Forewing with pterostigma wide. Vein Sc with stalk distant from R* making a kink to touch R and leaving Sc 1 oblique crossvein-like. 1-Rs present* single. 1-M shorter than 2-Cu. Cell 1mcu long* with its upper side three times as long as 1m-cu.

Aduantoma insolita Sun, Rasnitsyn, Zhuang & Shih gen. et sp. nov.

( Fig. 8) http://zoobank.org/ urn:lsid:zoobank.org:act:0041EF57-E06D-4083-A47D-26F6EC9D8DD3

Diagnosis: As for genus (see above).

Etymology: The species name is derived from the Latin word ‘insolitus’* meaning unusual* rare* and referring to the posture of this sawfly preserved gracefully.

Material: Holotype * CNU-HYM-NN2022310 p/c (part and counterpart).

Locality and horizon: Latest Middle Jurassic (late Callovian)* Jiulongshan Formation; Daohugou Village* Wuhua ºownship* Ningcheng County* Chifeng City* Inner Mongolia * China.

Description: Holotype * CNU-HYM-NN2022310 ( Fig. 8A–C). Male. Body length (excluding antennae) 8.4 mm. Forewing nearly complete* head* thorax* and part of legs preserved.

Head: ºransverse* somewhat narrowed forwards* with eyes occupying most of head sides* flat. Antennae not preserved.

Forewing: Length 6.5 mm as preserved. Pterostigma wide* sclerotized basally. Sc with stalk free (distant from R)* with kink to touch R at level of 1-M base* leaving Sc 1 as oblique crossvein intersecting with C behind level of 1-Rs ( Fig. 8D). 1-Rs distinct* single. 1-M (0.48 mm) 3.4 times as long as 1-Rs (0.14 mm)* gently arching* shorter than 2-Cu. Cell 1r large; Rs+M more than twice as long as 2-M. M+Cu bent. Crossvein 2r-rs (0.5 mm) nearly equal to 1r-rs (0.4 mm)* oblique* and meeting with pterostigma only near its apex. 1m-cu longer than 2-Cu.

Leg: Left hind leg well preserved. Protibia 2.0 mm long* with no spurs visible; tarsomere I longer than tarsomeres II* III* and IV combined; tarsomere lengths (in millimetres): 0.76* 0.32* 0.26* and 0.16* respectively; last segment not fully preserved.

Abdomen: Male genitalia ( Fig. 8E) with gonocoxa comparatively small* gonostylus long* penis long and wide.