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Order Grylloblattida Walker, 1914 Taxon Blattogryllopterida Cui, 2012 Remarks. This taxon and the Blattogryllidae as delimited by Storozhenko & Aristov (2014) have the same composition. Genus Plesioblattogryllus Huang, Nel & Petrulevičius, Published as part of CUI, YINGYING, BÉTHOUX, OLIVIER, YANG, NAN & REN, DONG, 2023, New data and systematic considerations on the Blattogryllopterida (Insecta: Grylloblattida) from the Jurassic Daohugou locality in China, pp. 124-132 in Palaeoentomology 6 (2) on page 125, DOI: 10.11646/palaeoentomology.6.2.4, http://zenodo.org/record/7929076, {"references":["Cui, Y. (2012) New data on the Blattogryllidae-Plesioblattogryllidae- Grylloblattidae complex (Insecta: Grylloblattida: Blattogryllopterida tax. n.). Arthropod Systematics & Phylogeny, 70 (3), 167 - 180.","Storozhenko & Aristov, D. S. (2014) Review of the Paleozoic and Mesozoic families Megakhosaridae and Blattogryllidae (Insecta: Grylloblattida). Far Eastern Entomologist, 271, 1 - 28.","Huang, D. Y., Nel, A. & Petrulevicius, J. F. (2008) New evolutionary evidence of Grylloblattida from the Middle Jurassic of Inner Mongolia, north-east China (Insecta, Polyneoptera). Zoological Journal of the Linnean Society, 152 (1), 17 - 24. https: // doi. org / 10.1111 / j. 1096 - 3642.2007.00351. x"]}

Prosyntexis lata Li, Shih & T. Gao sp. nov. (Figs 3–7) Type material. Holotype: CNU-HYM-MA2015108 (Figs 3, 4), female. Most of body preserved, except apex of right antenna lost; left antenna compressed diagenetically and right metatarsus fractured.The specimen is indistinct in left lateral view. Paratype: CNU-HYM-MA2015106 (Figs 5, 6), male. Most of body parts preserved except for missing apices of both antennae. CNU-HYM-MA2015109 (Fig. 7), male. Apices of forewings and hind legs missing. Etymology. The name lata is derived from the Latin word latus, referring to the broad pterostigma. Diagnosis. The new species is unique within Prosyntexis in having very long cell 1mcu (ratio of length to width more than 3). It also differs from other species, except P. gobiensis Rasnitsyn, 1993, in having 3r-m very close to 2m-cu in the forewing (ratio of length of 2m-cu to length of 3-M more than 5). Locality and horizon. Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. Holotype. Body about 6.52 mm long in lateral view, antenna 7.07 mm long in lateral view; forewing about 5.23 mm in length, maximum width 1.71 mm; hind wing about 4.01 mm in length (Fig. 3A, B). Head 0.57 mm long in lateral view. Compound eye medium sized and oval. Labial palpus with at least two palpomeres. Antenna elongate and filiform with 16 segments preserved; scape 0.23 mm long, maximum width 0.12 mm; pedicel 0.57 mm long, maximum width 0.09 mm; flagellomere I longer than scape and pedicel, 0.90 mm long, 0.10 mm wide, flagellomere II 0.66 mm long, 0.09 mm wide, as wide as remaining flagellomeres (Fig. 4B). Thorax with a deep medial membranous depression in the mesonotum bordered laterally by strong striate surfaces (Fig. 4A). Legs slender, profemur slightly longer than protibia; metafemur length 1.03 mm, metatibia 1.45 mm long, maximum width 0.13 mm. Single protibial spur, cleft apically (Fig. 4D); mesotibia with two apical spurs (Fig. 4E); metatibia with two apical spurs (the smaller spur less than half length of larger spur and not wide) (Fig. 4F). Tarsi pentamerous, basitarsomere long but shorter than remaining tarsomeres combined; tarsomere slightly elongate; pretarsal claws short, with curved apices and additional inner teeth (Fig. 4G). Forewing (Fig. 3C) with C strongly thickened; Sc+R present; costal space narrow, slightly enlarged at twothirds of its length; 1-Rs slanted toward wing apex and 0.8 as long as 1-M; pterostigma broad, oblong and 3.5× as long as wide; 1-M+Cu bent basally; distal part of 2- Rs undeveloped in both left and right forewings; 1r-rs connected to pterostigma at its base and slightly longer than 3-Rs; 2r-rs issuing from 1/3 of pterostigma and at nearly 0.4 of 2+3rm length; 2rs-m absent; 3-M extremely short and 0.2× as long as 2m-cu. Cell 1mcu subquadrate, about 2.6× as long as wide; cell 2mcu pentagonal, about 1.6× as long as wide. 2A+3A wavy medially. Hind wing (Fig. 3C) with cell r close; Sc+R present; 1-Rs about 0.99 mm in length, nearly 1.6× as long as 1-M; 1rs-m about 0.25 mm in length, short and slightly proclival; 1-M bent; 2-M+Cu present. 2rs-m, 3rs-m and 1m-cu absent; terminal abscissa of Cu and cu-a well-developed; 1-Cu and cu-a straight. Abdomen with short ovipositor sheath 1.85 mm long in ventral view and slightly protruding beyond apex of abdomen (Fig. 4C). Paratype (CNU-HYM-MA2015106). Body about 4.30 mm long in lateral view; forewing about 3.98 mm in length, maximum width 1.29 mm; hind wing about 3.02 mm in length (Figs 5, 6). Head 0.63 mm long and 0.43 mm wide in lateral view, rather small and subquadrate. Compound eyes medium sized and round. Maxillary palpus with six palpomeres (Fig. 6C); labial palpus with at least three palpomeres. Antenna elongate and filiform with 9 segments preserved; scape 0.20 mm long, maximum width 0.10 mm; pedicel 0.28 mm long, maximum width 0.10 mm, slightly swollen apically; flagellomere I longer than scape and pedicel, 0.56 mm long, 0.11 mm wide, flagellomere II 0.49 mm long, 0.11 mm wide (Fig. 6B). Thorax with deep medial membranous depression in the mesonotum bordered laterally by strong striate surfaces (Fig. 6D). Legs slender, profemur slightly longer than protibia; metafemur length 0.84 mm, metatibia 0.76 mm long, maximum width 0.14 mm. Protibia with one apical spur (Fig. 6F), mesotibia apical spurs not visible. Metatibia with two apical spurs (the smaller spur thick, less than half length of larger spur) (Fig. 6G). Tarsi pentamerous, basitarsomere long but shorter than remaining tarsomeres combined; metatarsomere elongate and claws 0.08 mm long with curved apices and additional inner teeth. Forewing (Fig. 5C) with C strongly thickened; Sc+R present; costal space distinctly narrow, slightly enlarged at the middle; basal section of Rs slanted toward wing apex and 1.6× as long as basal section of M; pterostigma broad, oblong and 2.8× as long as wide; 1-M+Cu bent basally; 2-M+Cu exceptionally short and nearly 0.05 mm long, scarcely present owing to proximal position of cua near origin of 1-Cu. 1-Rs+M 0.48 mm long and 4× as long as 2-Rs+M; end of 2-Rs very weak; 1r-rs connected to pterostigma at its base and longer than section of Rs between 1r-rs and 2r-rs; 2r-rs issuing from one-third of pterostigma and at one-third of 2+3rm length; 2rs-m absent; section of M between 2m-cu and 3rs-m extremely short and 0.2× as long as 2m-cu. Cell 1mcu subquadrate, about 3× as long as wide; cell 2mcu hexagonal, about 1.7× as long as wide. 2A+3A waved medially. Hind wing (Fig. 5C) with cell r closed and large; Sc+R present; 1-Rs about 0.86 mm in length, nearly 2× as long as 1-M; 1rs-m about 0.11 mm in length, short and slightly proclival; 1-M bent and 0.44 mm long; 2-M+Cu present and 0.20 mm long. 2rs-m, 3rs-m and 1m-cu absent; terminal abscissa of Cu and cu-a well-developed; 1-Cu and cu-a straight. Abdomen with nine distinguishable segments in lateral view. Hypopygium narrower rounded apically comparing P. aristovi sp. nov. Male genitalia (Fig. 6E) distinct, narrow, tightly closed obscuring details except gonostyle about 0.17 mm long, very narrow and with two thin threads of obscure identity at their apex. Cercus about 0.13 mm long. Paratype (CNU-HYM-MA2015109). Differs from the above paratype in shorter distance between compound eyes and postoccipital sulcus. Compound eyes rather large and oval.Antenna elongate and filiform with 16 segments, flagellomere I longer than scape and pedicel combined (Fig. 7A, B). Pronotum lateral edge slightly contracted inward. Hypopygium apparently as in other paratype. Genitalia internalized with very narrow gonostyli partially visible, with no threads visible., Published as part of LI, XIAO-QIN, RASNITSYN, ALEXANDR P., GAO, JIA, ZHANG, YAN-JIE, SHIH, CHUNG-KUN, REN, DONG, ZHAO, YUN-YUN & GAO, TAI-PING, 2023, New taxa of Sepulcidae (Hymenoptera) from mid-Cretaceous Kachin amber, pp. 133-145 in Palaeoentomology 6 (2) on pages 136-139, DOI: 10.11646/palaeoentomology.6.2.5, http://zenodo.org/record/7928967, {"references":["Rasnitsyn, A. P. (1993) New taxa of Sepulcidae. In: Ponomarenko, A. G. (Ed.), Mesozoic insects and ostracods from Asia. Transactions of the Paleontological Institute, Russian Academy of Sciences, 252, pp. 80 - 99. [In Russian]"]}

Key to species of Prosyntexis 1. Forewing with pterostigma narrow; 2r-rs joining pterostigma near its apex; cell 2r more than twice as long as wide................ P. montsecensis Rasnitsyn & Ansorge, 2000 - Forewing with pterostigma wide; 2r-rs joining pterostigma before its midlength; cell 2r less than twice as long as wide.................................................................................. 2 2. Forewing with costal area normal..................................................................................... P. gobiensis Rasnitsyn, 1993 - Forewing with basal third or half of costal area narrowed............................................................................................ 3 3. Forewing with cell 1mcu small, length to width ratio far less than 2........................... P. okhotensis Rasnitsyn, 1993 - Forewing with cell 1mcu normal size, length to width ratio more than 2....................................................................... 4 4. Forewing with 3r-m joining cell 3r almost at its midlength...................................... P. sennlaubi Jouault & Nel, 2021 - Forewing with 3r-m joining cell 3r at well before its midlength.......................................................................... 5 5. Forewing with 2-Rs not behind 1m-cu (no 2-RS+M); forewing length more than 10 mm......................................................................................... P. gouleti Sharkey, 1990 - Forewing with 2-Rs behind 1m-cu (2-RS+M present); forewing length less than 10 mm...................................... 6 6. Length of 2m-cu to the length of 3-M of forewing less than 2.................................. P. aristovi Li, Shih & Gao sp. nov. - Length of 2m-cu to the length of 3-M of forewing distinctly more than 2....................................................................... 7 7. Forewing with Sc present; 1-RS almost as long as 1-Rs+M; hind wing 2-M+Cu absent.................................................................................. P. antennata Li, Shih & Gao sp. nov. - Forewing with Sc absent; 1-RS short; hind wing 2-M+Cu present............................... P. lata Li, Shih & Gao sp. nov., Published as part of LI, XIAO-QIN, RASNITSYN, ALEXANDR P., GAO, JIA, ZHANG, YAN-JIE, SHIH, CHUNG-KUN, REN, DONG, ZHAO, YUN-YUN & GAO, TAI-PING, 2023, New taxa of Sepulcidae (Hymenoptera) from mid-Cretaceous Kachin amber, pp. 133-145 in Palaeoentomology 6 (2) on pages 142-143, DOI: 10.11646/palaeoentomology.6.2.5, http://zenodo.org/record/7928967, {"references":["Rasnitsyn, A. P. & Ansorge, J. (2000) New Early Cretaceous hymenopterous insects (Insecta: Hymenoptera) from Sierra del Montsec (Spain). Palaeontologische Zeitschrift, 74, 335 - 341. https: // doi. org / 10.1007 / BF 02988105","Rasnitsyn, A. P. (1993) New taxa of Sepulcidae. In: Ponomarenko, A. G. (Ed.), Mesozoic insects and ostracods from Asia. Transactions of the Paleontological Institute, Russian Academy of Sciences, 252, pp. 80 - 99. [In Russian]","Jouault, C. & Nel, A. (2021) A new species of Prosyntexis Sharkey, 1990 (Hymenoptera: Sepulcidae) from the Lower Cretaceous Crato Formation of Brazil confirmed by geometric morphometric analysis. Palaeoentomology, 4 (2), 171 - 177. https: // doi. org / 10.11646 / palaeoentomology. 4.2.6"]}

Prosyntexis antennata Li, Shih & T. Gao sp. nov. (Figs 8, 9) Type material. Holotype: CNU-HYM-MA2015107, a male specimen. Most of body preserved except left antenna broken and with apex lost, and left forewing broken. Etymology. The name antennata is derived from the Latin word antennatus referring to the long antennae. Diagnosis. The new species is unique within Prosyntexis in having cu-a behind the fork of M+Cu in the hind wing (hind wings not preserved or very incomplete in P. gouleti, P. gobiensis, P. montsecensis and P. aristovi). It also differs from other species, except for P. aristovi, in having a short and thin Sc in the forewing; and distinguished from P. aristovi by short 3-M (3r-m and 2m-cu approaching). Locality and horizon. Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. Body about 6.51 mm long in lateral view, antenna 8.31 mm long in lateral view; forewing about 5.89 mm in length, maximum width 2.56 mm; hind wing about 4.31 mm in length, maximum width 1.82 mm (Fig. 8A, B). Head 0.60 mm long and 0.68 mm height in lateral view, slightly small and subquadrate. Compound eyes slightly large and preserved in a shrunken state. In lateral view maxillary palpus with six palpomeres, palpomere I short and thick; palpomere II 0.33 mm long and swelled apically; palpomere III elongate and almost 0.71 mm long; palpomere IV 0.46 mm long; palpomere V 0.43 mm long and palpomere VI 0.44 mm. Labial palpus with two palpomeres with inflated last segment (Fig. 9C). Antenna elongate and filiform with 16 segments; scape 0.30 mm long, maximum width 0.12 mm; pedicel 0.42 mm long, maximum width 0.16 mm, slightly swollen apically; flagellomere I longer than scape and pedicel, 0.81 mm long, 0.15 mm wide, flagellomere II 0.74 mm long, 0.12 mm wide, as wide as individual widths of remaining flagellomeres (Fig. 9A). Thorax with membranous medial depression in the mesonotum bordered laterally by strong striate surfaces (Fig. 9D). Legs slender, profemur slightly longer than protibia; metafemur length 1.21 mm, metatibia 1.55 mm long, maximum width 0.22 mm. Single protibial spur, cleft apically (Fig. 9F); mesotibia with two apical spurs (Fig. 9G); metatibia with two apical spurs (smaller spur thick and less than half length of larger spur) (Fig. 9H). Tarsi pentamerous, basitarsomere long but shorter than remaining tarsomeres combined; tarsomere V not elongate; pretarsal claws short, with curved apices and additional inner teeth (Fig. 9I). Forewing (Fig. 8C) with C strongly thickened; costal space obviously narrow, slightly enlarged at middle; Sc short and thin, appearing like a crossvein before origin of Rs; 1-Rs slanted toward wing apex and twice as long as 1-M; pterostigma long and broad, oblong and 3.5× as long as wide with center translucent; M+Cu bent basally. 2-Rs+M especially short; junction of 2-Rs, 2-Rs+M and 2+3-M very weak; 1r-rs connected to pterostigma at its base and slightly shorter than section of Rs between 1r-rs and 2r-rs; 2r-rs issuing from 1/4 of pterostigma and at two-fifth of 2+3rm length; 2rs-m absent; 3-M short and 0.3× as long as 2m-cu. Cell 1mcu subquadrate, about 2.4× as long as wide; cell 2mcu hexagonal, about 2× as long as wide. 2A+3A wavy medially. Aberrant vein extending from 4-Cu near 2m-cu of left forewing, towards posterior margin of the wing (Fig. 8C). Hind wing (Fig. 8C) with basal and distal hamuli present (no intermediate ones found); cell r close; Sc+R present; 1-Rs about 1.08 mm in length, nearly 1.7× as long as 1-M; 1rs-m about 0.30 mm in length, proclival and short; 1-M bent; 2rs-m, 3rs-m and 1m-cu absent; terminal abscissa of Cu and cu-a well-developed; 1-Cu and cu-a straight, 1-Cu especially short and nearly 0.2× as long as cu-a in left hind wing; right hind wing lacking 1-Cu (cu-a coinciding with M+Cu fork). Abdomen with nine distinguishable segments in lateral view. Male genitalia (Fig. 9E) narrow, with gonostyle slender and about 0.36 mm long; cercus about 0.24 mm long., Published as part of LI, XIAO-QIN, RASNITSYN, ALEXANDR P., GAO, JIA, ZHANG, YAN-JIE, SHIH, CHUNG-KUN, REN, DONG, ZHAO, YUN-YUN & GAO, TAI-PING, 2023, New taxa of Sepulcidae (Hymenoptera) from mid-Cretaceous Kachin amber, pp. 133-145 in Palaeoentomology 6 (2) on pages 140-142, DOI: 10.11646/palaeoentomology.6.2.5, http://zenodo.org/record/7928967

Prosyntexis aristovi Li, Shih & T. Gao sp. nov. (Figs 1, 2) Type material. Holotype: CNU-HYM-MA2015105, male specimen. Most of body preserved, but major parts of head and mesonotum obscured by inclusions in ventral view. Left forewing and left and right hind wings folded over abdomen obscuring abdomen dorsally. Etymology. The specific name “ aristovi ” is dedicated to the memory of the late Dr Daniil Sergeevich Aristov, for his outstanding contribution to palaeoentomological research. Diagnosis. The new species is unique within Prosyntexis in having 2+3A widely interrupted before the cu-a crossvein. It also differs from other species in having a short and thin crossvein-like Sc in the forewing. Locality and horizon. Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. Body about 4.91 mm long, antenna 7.86 mm long in dorsal view; forewing about 4.66 mm in length, maximum width 2.08 mm (Fig 1A, B). Head nearly 0.93 mm wide and 0.67 mm long, rather large and subquadrate. Compound eyes rather small and oval. In ventral view, maxillary palpus with at least five palpomeres; labial palpus indiscernible. Antenna elongate and filiform with 16 segments; scape 0.22 mm long, maximum width 0.14 mm, slightly swollen apically; pedicel 0.40 mm long, maximum width 0.10 mm, slightly swollen apically and thinner than scape; flagellomere I longer than scape and pedicel 0.77 mm long, 0.10 mm wide; flagellomere II 0.76 mm long, 0.09 mm wide, as wide as individual widths of remaining flagellomeres (Fig. 2B). Thorax only partially visible with a deep medial membranous depression in the mesonotum bordered laterally by strong striate surfaces (Fig. 2C). Legs spindly, meso- and metafemur thicker than associated tibiae; metafemur length 0.95 mm, metatibia 1.50 mm long, maximum width 0.07 mm. Protibia with one apical spur (Fig. 2E), mesotibia with two apical spurs (Fig. 2F), metatibia with two apical spurs (the smaller spur thick, less than half length of larger spur) (Fig. 2G); basitarsomere long but shorter than remaining tarsomeres combined; tarsomere V not elongate; pretarsal claws short, with curved apices and additional inner teeth (Fig. 2D). Forewing (Fig. 1C) with C strongly thickened; R greatly thickened before base of Rs; basal third of costal space narrowed; crossvein-like Sc short and thin, appearing just before origin of Rs; 1-Rs slanted toward wing apex and 1.4× as long as 1-M; pterostigma broad, oblong and 3× as long as wide with center slightly translucent; 1-M+Cu bent basally; 2-M+Cu exceptionally short, nearly 0.05 mm long, scarcely present owing to the proximal position of cu-a near the origin of 1-Cu. 2- Rs+M short and 1/5× as long as 1-Rs+M; end of 2-Rs weakened; 1r-rs connected to pterostigma at its base and nearly as long as section of Rs between 1r-rs and 2r-rs; 2r-rs issuing from 1/4 of pterostigma and at 1/3 of 2+3rm length; 2rs-m absent; 3-M slightly shorter than 3rs-m. Cell 1mcu subquadrate, about 2× as long as wide; cell 2mcu hexagonal, about 1.5× as long as wide. 2+3A widely interrupted before 1a-2a. Hind wing with cell r closed, other parts indiscernible. Abdomen visible only ventrally, with hypopygium triangular with widely rounded apex; male genitalia not exerted, with only narrow gonocoxa partly visible., Published as part of LI, XIAO-QIN, RASNITSYN, ALEXANDR P., GAO, JIA, ZHANG, YAN-JIE, SHIH, CHUNG-KUN, REN, DONG, ZHAO, YUN-YUN & GAO, TAI-PING, 2023, New taxa of Sepulcidae (Hymenoptera) from mid-Cretaceous Kachin amber, pp. 133-145 in Palaeoentomology 6 (2) on page 134, DOI: 10.11646/palaeoentomology.6.2.5, http://zenodo.org/record/7928967

Aristoviblattogryllus minor (Ren & Aristov, 2011), comb. nov. (Figs 2, 3) Plesioblattogryllus minor Ren & Aristov, 2011: 36, fig. 1; Cui, 2012: 171, figs 2–5; Storozhenko & Aristov, 24. Diagnosis. The new genus is similar to Plesioblattogryllus but differs from latter in a long RP+MA fusion with a long basal free part of MA and a very short basal free section of RP. Material. Holotype CNU-GRY-NN2008001 and newly collected CNU-GRY-NN2023001. Type locality and horizon. Holotype and newly collected specimen are both from the Daohugou locality, Inner Mongolia, China; Middle Jurassic. Description. Mediumsizedinsects, bodylengthabout 30 mm, forewing length about 20–25 mm; head with large compound eyes and three ocelli; mandibles asymmetrical; pronotum rectangular with wide ring of paranotalia (lateral expansions of the pronotum); legs ambulatorial, at least fore tibia with a few long spines; coxa enlarged, fore and mid coxa rounded, hind coxa square-shaped; tarsi 5-segmented, segments 1–4 short with poorly preserved euplantulae, fifth segment elongate with a pair of claws and without arolia; female ovipositor straight, about 4.5 mm long; cerci with 10–11 segments, about 7.5 mm long; eggs olive-shaped with strong longitudinal ridges. The details of wing-venation variability are given below. Holotype CNU-GRY-NN2008001 (Fig. 2): positive and negative imprints of left wing, likely a forewing, with distal part of AA area missing; 24.6 mm long, 7.4 mm wide; ScP ending on anterior wing margin; in distal portion of ScP–RA area crossveins desclerotized along line parallel to ScP and RA, located roughly in middle of area (likely indicating occurrence of flexion line); RA/RP fork in basal third of wing length; RA simple, nearly parallel to ScP; area between anterior wing margin and ScP slightly narrower than ScP–RA area; RP posteriorly pectinate, with four branches; RA–RP area very narrow, parallel to areas between anterior wing margin and ScP, and ScP–RA area; bases of M and CuA free for short distance; CuA arising early, then fused with M for 3.0 mm; MA diverging from M+CuA near basal fourth of wing length, then fused with RP for 3.1 mm, slightly after origin of RP; MA simple; MP diverging from MP+CuA slightly after origin of MA; MP simple, with middle part desclerotised; CuA forked into CuA1 and CuA2 1.2 mm after its origin from MP+CuA; CuA1/CuA2 fork nearly opposite to RP/MA fork; CuA2 simple; CuP strong, straight; AA with anterior-most vein forked; no evident ppa, possibly located basal to second AA vein; cross-veins scalariform, rarely reticulated. Specimen CNU-GRY-NN2023001 (Fig. 3): negative imprint of right wing, presumably hind wing, with posterior part of anal area folded; 19.6 mm long, 7.2 mm wide as preserved; ScP ending on anterior wing margin; in distal portion of ScP–RA area crossveins desclerotized; RA/RP fork slightly after basal fourth of wing length; RA simple and parallel to ScP; RP posteriorly pectinate, with five branches; area between anterior wing margin and ScP, and RA–RP area slightly narrower than ScP–RA area; base of M invisible; MA diverges from M+CuA in basal sixth of wing length; MA fused with RP for 2.5 long slightly after origin of RP; MA simple; MP diverges from MP+CuA slightly after origin of MA; MP simple, with middle part desclerotised; CuA forks into CuA1 and CuA2 slightly after its origin from MP+CuA; CuA forked earlier than divergence of MA from RP+MA; CuA2 simple; CuP strong, straight; AA with anteriormost vein forked, and both branches crossed by fold (presumably ppa); crossveins scalariform, rarely reticulated. Remarks. The new specimen CNU-GRY-NN2023001 is roughly identical to the holotype of A. minor except for a slightly smaller size, a 5-branched RP (4-branched in the holotype) and a possibly slightly expanded plicatum. In contrast with the comparatively larger size of the holotype, with 25 mm in wing length, the multiple specimens investigated by Cui (2012) indicated an average wing length for A. minor around 20 mm, including female individuals, which is consistent with an assignment of the new specimen to this species. However, the new specimen is also wider than the holotype. As a consequence, both specimens differ in general shape. Plastic deformation, already assumed for some specimens from the same locality (Cui et al., 2015: figs 7C, 8D), may have affected the specimen and caused the observed shape variation. A moderate elongation affecting the holotype along its longitudinal axis, and a moderate elongation affecting the new specimen along the transverse axis, could explain the observed difference. Thus, here we consider that the size/ shape difference displayed by the new material is under the range of intra-specific variation. In contrast with the general perception that a smaller size entails a reduced venation, the new material exhibits a 5-branched RP, while this vein has 4 branches in most specimens of A. minor, including the holotype. The range of variation of this character is not well-documented for Blattogryllopterida. However, at the level of the order Grylloblattida, such variation commonly occurs within a single species, as demonstrated by species known from large numbers of specimens, and specimens preserving complete wing pairs (Storozhenko, 1998; Béthoux & Nel, 2010; Aristov & Mostovski, 2013; Storozhenko & Aristov, 2014; Cui et al., 2015, 2022, and others). Thus, a variation ranging from 4 to 5 branches for RP is very likely in A. minor. Our observation of the new specimen shows that the two branches of the first anal vein are folded along a straight line (orange dashed line on Fig. 4B). Two possible scenarios on the interpretation of this line are proposed, namely (i) an accidental fold affecting a forewing; and (ii) the occurrence of a plica prima anterior (ppa) crossing AA veins, suggestive of a hind wing. The second scenario is plausible, since ppa is commonly oblique in hind wings of polyneopteran insects. And although it rarely crosses veins in hind wing (but see Béthoux et al., 2011: fig. 3), such configuration cannot be excluded. Notably, in forewings of Mantodea in which the ppa is secondarily oblique, resulting in a somewhat enlarged plicatum, it crosses veins (Fig. 4; and see Demers-Potvin et al., 2021; Béthoux & Wieland, 2009). Under this scenario, the hind wing plicatum would be distinctly reduced in A. minor (but slightly enlarged compared to the forewing yet; Fig. 2B). Up to now, this state has never been documented in Grylloblattida (Storozhenko, 1998). It is worth mentioning that, within Polyneoptera, a secondarily reduced hind wing plicatum has been acquired in Embioptera and within Isoptera (Ross, 2000; Engel et al., 2009). A third case of reduction is therefore plausible. If this is correct, possessing a reduced hind wing plicatum may compose a distinctive trait of a yet unnamed taxon within the Blattogryllopterida. Therefore, all the observed differences between the new material and the holotype of A. minor are considered as intra-specific variation.Further substantial investigation on wing venation, especially the hind wing, is needed.

Prosyntexis aristovi Li, Shih & T. Gao sp. nov. (Figs 1, 2) Type material. Holotype: CNU-HYM-MA2015105, male specimen. Most of body preserved, but major parts of head and mesonotum obscured by inclusions in ventral view. Left forewing and left and right hind wings folded over abdomen obscuring abdomen dorsally. Etymology. The specific name “ aristovi ” is dedicated to the memory of the late Dr Daniil Sergeevich Aristov, for his outstanding contribution to palaeoentomological research. Diagnosis. The new species is unique within Prosyntexis in having 2+3A widely interrupted before the cu-a crossvein. It also differs from other species in having a short and thin crossvein-like Sc in the forewing. Locality and horizon. Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. Body about 4.91 mm long, antenna 7.86 mm long in dorsal view; forewing about 4.66 mm in length, maximum width 2.08 mm (Fig 1A, B). Head nearly 0.93 mm wide and 0.67 mm long, rather large and subquadrate. Compound eyes rather small and oval. In ventral view, maxillary palpus with at least five palpomeres; labial palpus indiscernible. Antenna elongate and filiform with 16 segments; scape 0.22 mm long, maximum width 0.14 mm, slightly swollen apically; pedicel 0.40 mm long, maximum width 0.10 mm, slightly swollen apically and thinner than scape; flagellomere I longer than scape and pedicel 0.77 mm long, 0.10 mm wide; flagellomere II 0.76 mm long, 0.09 mm wide, as wide as individual widths of remaining flagellomeres (Fig. 2B). Thorax only partially visible with a deep medial membranous depression in the mesonotum bordered laterally by strong striate surfaces (Fig. 2C). Legs spindly, meso- and metafemur thicker than associated tibiae; metafemur length 0.95 mm, metatibia 1.50 mm long, maximum width 0.07 mm. Protibia with one apical spur (Fig. 2E), mesotibia with two apical spurs (Fig. 2F), metatibia with two apical spurs (the smaller spur thick, less than half length of larger spur) (Fig. 2G); basitarsomere long but shorter than remaining tarsomeres combined; tarsomere V not elongate; pretarsal claws short, with curved apices and additional inner teeth (Fig. 2D). Forewing (Fig. 1C) with C strongly thickened; R greatly thickened before base of Rs; basal third of costal space narrowed; crossvein-like Sc short and thin, appearing just before origin of Rs; 1-Rs slanted toward wing apex and 1.4× as long as 1-M; pterostigma broad, oblong and 3× as long as wide with center slightly translucent; 1-M+Cu bent basally; 2-M+Cu exceptionally short, nearly 0.05 mm long, scarcely present owing to the proximal position of cu-a near the origin of 1-Cu. 2- Rs+M short and 1/5× as long as 1-Rs+M; end of 2-Rs weakened; 1r-rs connected to pterostigma at its base and nearly as long as section of Rs between 1r-rs and 2r-rs; 2r-rs issuing from 1/4 of pterostigma and at 1/3 of 2+3rm length; 2rs-m absent; 3-M slightly shorter than 3rs-m. Cell 1mcu subquadrate, about 2× as long as wide; cell 2mcu hexagonal, about 1.5× as long as wide. 2+3A widely interrupted before 1a-2a. Hind wing with cell r closed, other parts indiscernible. Abdomen visible only ventrally, with hypopygium triangular with widely rounded apex; male genitalia not exerted, with only narrow gonocoxa partly visible.

Aristoviblattogryllus minor (Ren & Aristov, 2011), comb. nov. (Figs 2, 3) Plesioblattogryllus minor Ren & Aristov, 2011: 36, fig. 1; Cui, 2012: 171, figs 2–5; Storozhenko & Aristov, 24. Diagnosis. The new genus is similar to Plesioblattogryllus but differs from latter in a long RP+MA fusion with a long basal free part of MA and a very short basal free section of RP. Material. Holotype CNU-GRY-NN2008001 and newly collected CNU-GRY-NN2023001. Type locality and horizon. Holotype and newly collected specimen are both from the Daohugou locality, Inner Mongolia, China; Middle Jurassic. Description. Mediumsizedinsects, bodylengthabout 30 mm, forewing length about 20–25 mm; head with large compound eyes and three ocelli; mandibles asymmetrical; pronotum rectangular with wide ring of paranotalia (lateral expansions of the pronotum); legs ambulatorial, at least fore tibia with a few long spines; coxa enlarged, fore and mid coxa rounded, hind coxa square-shaped; tarsi 5-segmented, segments 1–4 short with poorly preserved euplantulae, fifth segment elongate with a pair of claws and without arolia; female ovipositor straight, about 4.5 mm long; cerci with 10–11 segments, about 7.5 mm long; eggs olive-shaped with strong longitudinal ridges. The details of wing-venation variability are given below. Holotype CNU-GRY-NN2008001 (Fig. 2): positive and negative imprints of left wing, likely a forewing, with distal part of AA area missing; 24.6 mm long, 7.4 mm wide; ScP ending on anterior wing margin; in distal portion of ScP–RA area crossveins desclerotized along line parallel to ScP and RA, located roughly in middle of area (likely indicating occurrence of flexion line); RA/RP fork in basal third of wing length; RA simple, nearly parallel to ScP; area between anterior wing margin and ScP slightly narrower than ScP–RA area; RP posteriorly pectinate, with four branches; RA–RP area very narrow, parallel to areas between anterior wing margin and ScP, and ScP–RA area; bases of M and CuA free for short distance; CuA arising early, then fused with M for 3.0 mm; MA diverging from M+CuA near basal fourth of wing length, then fused with RP for 3.1 mm, slightly after origin of RP; MA simple; MP diverging from MP+CuA slightly after origin of MA; MP simple, with middle part desclerotised; CuA forked into CuA1 and CuA2 1.2 mm after its origin from MP+CuA; CuA1/CuA2 fork nearly opposite to RP/MA fork; CuA2 simple; CuP strong, straight; AA with anterior-most vein forked; no evident ppa, possibly located basal to second AA vein; cross-veins scalariform, rarely reticulated. Specimen CNU-GRY-NN2023001 (Fig. 3): negative imprint of right wing, presumably hind wing, with posterior part of anal area folded; 19.6 mm long, 7.2 mm wide as preserved; ScP ending on anterior wing margin; in distal portion of ScP–RA area crossveins desclerotized; RA/RP fork slightly after basal fourth of wing length; RA simple and parallel to ScP; RP posteriorly pectinate, with five branches; area between anterior wing margin and ScP, and RA–RP area slightly narrower than ScP–RA area; base of M invisible; MA diverges from M+CuA in basal sixth of wing length; MA fused with RP for 2.5 long slightly after origin of RP; MA simple; MP diverges from MP+CuA slightly after origin of MA; MP simple, with middle part desclerotised; CuA forks into CuA1 and CuA2 slightly after its origin from MP+CuA; CuA forked earlier than divergence of MA from RP+MA; CuA2 simple; CuP strong, straight; AA with anteriormost vein forked, and both branches crossed by fold (presumably ppa); crossveins scalariform, rarely reticulated. Remarks. The new specimen CNU-GRY-NN2023001 is roughly identical to the holotype of A. minor except for a slightly smaller size, a 5-branched RP (4-branched in the holotype) and a possibly slightly expanded plicatum. In contrast with the comparatively larger size of the holotype, with 25 mm in wing length, the multiple specimens investigated by Cui (2012) indicated an average wing length for A. minor around 20 mm, including female individuals, which is consistent with an assignment of the new specimen to this species. However, the new specimen is also wider than the holotype. As a consequence, both specimens differ in general shape. Plastic deformation, already assumed for some specimens from the same locality (Cui et al., 2015: figs 7C, 8D), may have affected the specimen and caused the observed shape variation. A moderate elongation affecting the holotype along its longitudinal axis, and a moderate elongation affecting the new specimen along the transverse axis, could explain the observed difference. Thus, here we consider that the size/ shape difference displayed by the new material is under the range of intra-specific variation. In contrast with the general perception that a smaller size entails a reduced venation, the new material exhibits a 5-branched RP, while this vein has 4 branches in most specimens of A. minor, including the holotype. The range of variation of this character is not well-documented for Blattogryllopterida. However, at the level of the order Grylloblattida, such variation commonly occurs within a single species, as demonstrated by species known from large numbers of specimens, and specimens preserving complete wing pairs (Storozhenko, 1998; Béthoux & Nel, 2010; Aristov & Mostovski, 2013; Storozhenko & Aristov, 2014; Cui et al., 2015, 2022, and others). Thus, a variation ranging from 4 to 5 branches for RP is very likely in A. minor. Our observation of the new specimen shows that the two branches of the first anal vein are folded along a straight line (orange dashed line on Fig. 4B). Two possible scenarios on the interpretation of this line are proposed, namely (i) an accidental fold affecting a forewing; and (ii) the occurrence of a plica prima anterior (ppa) crossing AA veins, suggestive of a hind wing. The second scenario is plausible, since ppa is commonly oblique in hind wings of polyneopteran insects. And although it rarely crosses veins in hind wing (but see Béthoux et al., 2011: fig. 3), such configuration cannot be excluded. Notably, in forewings of Mantodea in which the ppa is secondarily oblique, resulting in a somewhat enlarged plicatum, it crosses veins (Fig. 4; and see Demers-Potvin et al., 2021; Béthoux & Wieland, 2009). Under this scenario, the hind wing plicatum would be distinctly reduced in A. minor (but slightly enlarged compared to the forewing yet; Fig. 2B). Up to now, this state has never been documented in Grylloblattida (Storozhenko, 1998). It is worth mentioning that, within Polyneoptera, a secondarily reduced hind wing plicatum has been acquired in Embioptera and within Isoptera (Ross, 2000; Engel et al., 2009). A third case of reduction is therefore plausible. If this is correct, possessing a reduced hind wing plicatum may compose a distinctive trait of a yet unnamed taxon within the Blattogryllopterida. Therefore, all the observed differences between the new material and the holotype of A. minor are considered as intra-specific variation.Further substantial investigation on wing venation, especially the hind wing, is needed., Published as part of CUI, YINGYING, BÉTHOUX, OLIVIER, YANG, NAN & REN, DONG, 2023, New data and systematic considerations on the Blattogryllopterida (Insecta: Grylloblattida) from the Jurassic Daohugou locality in China, pp. 124-132 in Palaeoentomology 6 (2) on pages 127-131, DOI: 10.11646/palaeoentomology.6.2.4, http://zenodo.org/record/7929076, {"references":["Ren, D. & Aristov, D. S. (2011) New species of Plesioblattogryllus Huang, Nel et Petrulevicius (Grylloblattida: Plesioblattogryl- lidae) from the Middle Jurassic of China. Paleontologicheskii Zhurnal, 2011 (3), 36 - 37. [In Russian; English translation: Paleontological Journal, 45 (3), 273 - 274.] https: // doi. org / 10.1134 / S 0031030111030178","Cui, Y. (2012) New data on the Blattogryllidae-Plesioblattogryllidae- Grylloblattidae complex (Insecta: Grylloblattida: Blattogryllopterida tax. n.). Arthropod Systematics & Phylogeny, 70 (3), 167 - 180.","Cui, Y., Bethoux, O., Klass, K. - D. & Ren, D. (2015) The Jurassic Bajanzhargalanidae (Insecta: Grylloblattida?): new genera and species, and data on postabdominal morphology. Arthropod Structure & Development, 44 (6), 688 - 716. https: // doi. org / 10.1016 / j. asd. 2015.04.008","Storozhenko, S. Y. (1998) Systematics, phylogeny, and evolution of grylloblattids (Insecta: Grylloblattida). Dalnauka, Vladivostok, 207 pp. [In Russian]","Bethoux, O. & Nel, A. (2010) Description of a new grylloblattidan insect from Montceau-les-Mines (Late Carboniferous; France) and definition of Phenopterum Carpenter, 1950. Systematic Entomology, 35 (3), 546 - 553. https: // doi. org / 10.1111 / j. 1365 - 3113.2010.00527. x","Aristov, D. S. & Mostovski, M. B. (2013) Grylloblattids (Insecta, Grylloblattida) from the Upper Permian of South Africa. Paleontologicheskii Zhurnal, 5, 37 - 43. [In Russian; English translation: Paleontological Journal, 47 (5), 495 - 502.] https: // doi. org / 10.1134 / S 0031030113050031","Storozhenko & Aristov, D. S. (2014) Review of the Paleozoic and Mesozoic families Megakhosaridae and Blattogryllidae (Insecta: Grylloblattida). Far Eastern Entomologist, 271, 1 - 28.","Cui, Y., Brauner, S., Schneider, J. W. & Bethoux, O. (2022) Grylloblattidan insects from Sperbersbach and Cabarz (Germany), two new early Permian and insect-rich localities. Journal of Paleontology, 96 (2), 355 - 374. https: // doi. org / 10.1017 / jpa. 2021.101","Bethoux, O., Schneider, J. W. & Klass, K. - D. (2011) Redescription of the holotype of Phyloblatta gaudryi (Agnus, 1903) (Pennsylvanian; Commentry, France), an exceptionally well-preserved stem-dictyopteran. Geodiversitas, 33 (4), 625 - 635. https: // doi. org / 10.5252 / g 2011 n 4 a 4","Demers-Potvin, A. V., Larsson, H. C. E., Cournoyer, M. & Bethoux, O. (2021) Wing morphology of a new Cretaceous praying mantis solves the phylogenetic jigsaw of early-diverging extant lineages. Systematic Entomology, 46 (1), 205 - 223. https: // doi. org / 10.1111 / syen. 12457","Bethoux, O. & Wieland, F. (2009) Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology. Zoological Journal of the Linnean Society, 156 (1), 79 - 113. https: // doi. org / 10.1111 / j. 1096 - 3642.2008.00485. x","Ross, E. A. (2000) Contributions to the biosystematics of the insect order Embiidina. Part 1. Origin, relationships and integumental anatomy of the insect order Embiidina. Occasional papers of the California Academy of Sciences, 149, 1 - 53.","Engel, M. S., Grimaldi, D. & Krishna, K. (2009) Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance. American Museum Novitates, 3650, 1 - 27. https: // doi. org / 10.1206 / 651.1"]}

Prosyntexis antennata Li, Shih & T. Gao sp. nov. (Figs 8, 9) Type material. Holotype: CNU-HYM-MA2015107, a male specimen. Most of body preserved except left antenna broken and with apex lost, and left forewing broken. Etymology. The name antennata is derived from the Latin word antennatus referring to the long antennae. Diagnosis. The new species is unique within Prosyntexis in having cu-a behind the fork of M+Cu in the hind wing (hind wings not preserved or very incomplete in P. gouleti, P. gobiensis, P. montsecensis and P. aristovi). It also differs from other species, except for P. aristovi, in having a short and thin Sc in the forewing; and distinguished from P. aristovi by short 3-M (3r-m and 2m-cu approaching). Locality and horizon. Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, mid-Cretaceous. Description. Body about 6.51 mm long in lateral view, antenna 8.31 mm long in lateral view; forewing about 5.89 mm in length, maximum width 2.56 mm; hind wing about 4.31 mm in length, maximum width 1.82 mm (Fig. 8A, B). Head 0.60 mm long and 0.68 mm height in lateral view, slightly small and subquadrate. Compound eyes slightly large and preserved in a shrunken state. In lateral view maxillary palpus with six palpomeres, palpomere I short and thick; palpomere II 0.33 mm long and swelled apically; palpomere III elongate and almost 0.71 mm long; palpomere IV 0.46 mm long; palpomere V 0.43 mm long and palpomere VI 0.44 mm. Labial palpus with two palpomeres with inflated last segment (Fig. 9C). Antenna elongate and filiform with 16 segments; scape 0.30 mm long, maximum width 0.12 mm; pedicel 0.42 mm long, maximum width 0.16 mm, slightly swollen apically; flagellomere I longer than scape and pedicel, 0.81 mm long, 0.15 mm wide, flagellomere II 0.74 mm long, 0.12 mm wide, as wide as individual widths of remaining flagellomeres (Fig. 9A). Thorax with membranous medial depression in the mesonotum bordered laterally by strong striate surfaces (Fig. 9D). Legs slender, profemur slightly longer than protibia; metafemur length 1.21 mm, metatibia 1.55 mm long, maximum width 0.22 mm. Single protibial spur, cleft apically (Fig. 9F); mesotibia with two apical spurs (Fig. 9G); metatibia with two apical spurs (smaller spur thick and less than half length of larger spur) (Fig. 9H). Tarsi pentamerous, basitarsomere long but shorter than remaining tarsomeres combined; tarsomere V not elongate; pretarsal claws short, with curved apices and additional inner teeth (Fig. 9I). Forewing (Fig. 8C) with C strongly thickened; costal space obviously narrow, slightly enlarged at middle; Sc short and thin, appearing like a crossvein before origin of Rs; 1-Rs slanted toward wing apex and twice as long as 1-M; pterostigma long and broad, oblong and 3.5× as long as wide with center translucent; M+Cu bent basally. 2-Rs+M especially short; junction of 2-Rs, 2-Rs+M and 2+3-M very weak; 1r-rs connected to pterostigma at its base and slightly shorter than section of Rs between 1r-rs and 2r-rs; 2r-rs issuing from 1/4 of pterostigma and at two-fifth of 2+3rm length; 2rs-m absent; 3-M short and 0.3× as long as 2m-cu. Cell 1mcu subquadrate, about 2.4× as long as wide; cell 2mcu hexagonal, about 2× as long as wide. 2A+3A wavy medially. Aberrant vein extending from 4-Cu near 2m-cu of left forewing, towards posterior margin of the wing (Fig. 8C). Hind wing (Fig. 8C) with basal and distal hamuli present (no intermediate ones found); cell r close; Sc+R present; 1-Rs about 1.08 mm in length, nearly 1.7× as long as 1-M; 1rs-m about 0.30 mm in length, proclival and short; 1-M bent; 2rs-m, 3rs-m and 1m-cu absent; terminal abscissa of Cu and cu-a well-developed; 1-Cu and cu-a straight, 1-Cu especially short and nearly 0.2× as long as cu-a in left hind wing; right hind wing lacking 1-Cu (cu-a coinciding with M+Cu fork). Abdomen with nine distinguishable segments in lateral view. Male genitalia (Fig. 9E) narrow, with gonostyle slender and about 0.36 mm long; cercus about 0.24 mm long.

Burmotachinymphes pengi Chen, Deng et Yang sp. nov. (Fig. 1) urn:lsid:zoobank.org:act: BD3B288A-383D-4A64-8AA7-7F4D6B657480 Material. Holotype CNU-NEU-MA2018075, preserved nearly complete left wings (wing base not preserved). Type locality and horizon. Hukawng Valley of the northern state of Kachin in Myanmar (26°200’N, 96°360’E); lowermost Cenomanian, Upper Cretaceous. Etymology. The specific name is after Mr Wei Peng (Jilin, China), who is very interested in Entomology and donated the specimen for this study. Diagnosis. Forewing RP with thirteen branches, MP with six distal pectinate branches, CuA with five distal pectinate branches. Hind wing RP with ten branches, MP with five distal pectinate branches, CuA with about seven distal pectinate branches. Conspicuous pterostigma present in hind wing (not preserved in forewing). Pterostigmal area with eight veinlets. Description. Forewing: Length about 18.3 mm (complete length estimated more than 20 mm), width about 6.6 mm as preserved. Forewing relatively broad, with rounded apex. Trichosors absent. Costal space straight, with simple unforked veinlets, equally narrow. Pterostigma not preserved. Endings of ScP and RA indistinct. Subcostal space broad, no crossvein detected. The space between RA and RP broad in proximal part, narrowed towards wing apex. Thirteen crossveins detected between RA and RP. Stem of RP smooth and zigzagged, with thirteen regularly zigzagged branches. Numerous crossveins present in radial space, venation reticulated, gradate series inconspicuous, inner gradate series better defined than outer gradate series. Crossvein present between MA and RP before the origin of RP1, connecting stem of RP and intramedian cell. Basal part of M and its dividing into MA and MP not preserved. MA simple. MP with six pectinate terminal branches. CuA with five pectinate terminal branches. RP, MA, MP and CuA not coalesced. CuP poorly preserved. A not preserved. Hind wing: Length about 16.3 mm, width about 5.2 mm as preserved. Trichosors absent. Costal space similar with forewing. Pterostigma conspicuous. Pterostigma area with eight veinlets. ScP fused with RA apically, then end on costal margin near wing apex. Subcostal space relatively narrow, crossveins not detected. The space between RA and RP broad in proximal part, narrowed towards wing apex, with eleven crossveins. Stem of RP smooth and zigzagged, with ten regularly, zigzagged branches. Numerous crossveins in radial space present, venation reticulated, gradate series inconspicuous, inner gradate series better defined than outer gradate series. Crossvein present between MA and RP before the origin of RP1, connecting stem of RP and base of second intramedian cell. M divided into MA and MP before the origin of RP1. MA simple. MP with five pectinate terminal branches. CuA with about seven pectinate terminal branches. RP, MA, MP and CuA not coalesced. A not preserved. Remarks. The new species is assigned to Burmotachinymphes based on pterostigma obvious; costal crossveins simple; RP diverged from RA at an angle of>30°; RP with zigzagged stem; crossveins in RP area forming gradate series; MA simple, MP with pectinate branches. It is easy to distinguish from Burmotachinymphes bilobata by body size. The preserved part of forewing of Burmotachinymphes pengi sp. nov. is more than 18 mm in length, and the estimated complete length more than 20 mm. The hind wing is more 16 mm in length. In contrast, Burmotachinymphes bilobata has forewing long 12.7 mm by estimation, hind wing long 10.9 mm. In addition, there are some other distinct differences between them: Burmotachinymphes bilobata RP has seven branches with RP1 to RP3 each bearing two marginal branches and RP4 to RP7 simple in fore- and hind wings. MP has five distal marginal pectinate branches in forewing, and four in hind wing. CuA has four distal marginal pectinate branches in fore- and hind wings. The new species has thirteen RP branches with RP1 to RP8 each bearing at least two marginal branches and RP9 to RP13 simple in forewing. In hind wing, there are ten RP branches with RP1 to RP6 each bearing two marginal branches and RP7 to RP10 simple. MP has six distal marginal pectinate branches in forewing, and five distal marginal pectinate branches in hind wing. CuA has five distal marginal pectinate branches in forewing and about seven distal marginal pectinate branches in hind wing. Tachinymphes is the largest genus in this subfamily with six species. Nanochrysopa Nel et al., 2005 is a monotypic genus. Burmotachinymphes shares characters with other genera of Tachinymphinae in the MP not touching CuA. Besides, Burmotachinymphes have several unique characters, i.e., 1) MA simple, 2) MP and CuA with pectinate distal marginal branches, and 3) RP, MA, MP and CuA not coalesced evidently., Published as part of Chen, Xinrui, Chen, Peichao, Deng, Xinru, Yang, Qiang, Shi, Chaofan & Ren, Dong, 2022, A new mesochrysopid (Insecta: Neuroptera) from Upper Cretaceous, pp. 597-600 in Zootaxa 5134 (4) on pages 598-599, DOI: 10.11646/zootaxa.5134.4.8, http://zenodo.org/record/6541823

Panorpidium validum sp. nov. (Fig. 3) urn:lsid:zoobank.org:act: 58900EF2-F7E7-421E-B840-07B5D975D7C2 Diagnosis. Forewing medium sized, the end of ScA close to the end of CP, R forking into RA and RP at the level of ScP reaching anterior wing margin, CuPa subequal to CuPaα in length. Material. Holotype, CNU-ORT-LB2016067, female. Etymology. From the latin ‘ validus ’ for strong, refers to its unusually strong hind femur. Horizon and locality. Yixian Formation, Lower Cretaceous, Huangbanjigou Village, Beipiao City, Liaoning Province, China. General Description. A well-preserved female, the body medium size, head preserved much poorly and abdomen was been compressed. Head. Compound eye large, suborbicular. Thorax. Pronotum saddle-shaped with post margin prolongation. Prothorax leg preserved fragmentarily. Basal and middle part of metafemur much strong and distal part rapidly narrowed, inner side of it with irregular dark spots; hind knee robust, semi-lunar processes prominent on the dorsal edge; metatibia long and slender, dorsum with a row of tiny spines in basal half and two pairs of big leaf-like spurs in distal half visible; leaf-like spurs narrow, long and thickened, with a rachis in the middle part, then gradually thinned to the margin; basitarsomere slender, only preserved very basal part. Abdomen. Abdomen short, probably about half the length of the wing. Ovipositor gently recurved and elongated, its length more than two times than pronotum; the surface of the vavulae smooth. Wings. Forewing size large, Costa area relatively broad, CP reaching anterior margin about 1/3 length of forewing, with numerous branchlets; ScA reaching anterior margin at about origin of MP, with 3 branchlets connect with CP and distally simple; area between ScA and CP narrower than that of anterior and CP; ScP ending in anterior margin at about 1/2 length of forewing, several short branches connecting with ScA and four branches distally reaching anterior margin; stem R long and distinctly strong, branched into RA and RP near the ending of ScP; area between R and ScP relatively broad; RA ending in anterior margin close to apex and producing numerous oblique branches; RP with 11 comb-like branches, most of them reaching posterior margin, several distal terminals reaching anterior margin; most cross-veins between the branches of RP simple and straight; area between RA and RP narrow; M forking into MA and MP at about terminals of ScA; MA branching into MA1 and MA2 close to the last branch point of ScP; MA1 with 2 branches, with the first fused with RP and then forked immediately; MA2 simple; MP simple, originating at about terminal of ScA and reaching posterior margin at about 3/4 distal of forewing base; CuA+CuPaα simple, starting before the end point of CP, ending in posterior margin at about 2/3 length of forewing; CuPa straight, slanted toward the anterior margin, nearly as long as CuPaα; CuPaβ, CuPb and AA1 fused with each other at about 1/3rd of wing length; area between CuPb and AA1 relatively narrow, AA1 straight and strong; AA2 straight; area between AA1 and posterior margin filled with simple and straight cross-veins. Area between ScP and R, cross-veins between RA and RP, and that of CuPaβ, CuPb and AA1 are colored, one round dark spots located at the branching point of CuPa. Measurement. The specimen preserved very well but missing most part of head. The body total estimated length approximately 11 mm (from head to the end of abdomen without cercus and ovipositor). Pronotum 3.2 mm long. Left profemur preserved 2.2 mm long, protibia preserved 1.9 mm long. Left metathorax leg: metafemur preserved 5.6 mm long, metatibia 9.9 mm long and 0.5 mm wide, ds2 preserved 1.5 mm long and 0.4 mm wide, ds3 2.1 mm long and 0.4 mm wide, basitarsomere preserved 1.4 mm long. Right metathorax leg: metafemur preserved 7.7 mm long, metatibia 10.2 mm long and 0.5 mm wide, ds1 1.8 mm long, ds2 preserved 2.3 mm long and 0.4 mm wide. Ovipositor preserved 7.6 mm long. Left forewing 20.5 mm long and 4.5 mm wide (the maximum width). Remarks. This new species can be assigned to Elcaninae by its relatively narrow area between RA and RP, and CuPaβ, CuPb, and AA1 fused with each other. It falls in Panorpidium Westwood, 1854 by few branched ScA; three longitudinal branches between basal part of RP and CuA+CuPaα; and CuPaβ, CuPb and AA1 fused with each other distally (Fig. 3D–E). This new species is distinguished from all species of Panorpidium in the end of ScA close to the end of CP, CuPa straight and subequal to CuPaα in length and different pattern of coloration., Published as part of Tian, He, Ren, Dong & Gu, Jun-Jie, 2022, Two new species of Elcaninae (Orthoptera, Elcanidae) recorded from the Early Cretaceous China, pp. 589-596 in Zootaxa 5128 (4) on page 593, DOI: 10.11646/zootaxa.5128.4.8, http://zenodo.org/record/6480106

Panorpidium validum sp. nov. (Fig. 3) urn:lsid:zoobank.org:act: 58900EF2-F7E7-421E-B840-07B5D975D7C2 Diagnosis. Forewing medium sized, the end of ScA close to the end of CP, R forking into RA and RP at the level of ScP reaching anterior wing margin, CuPa subequal to CuPaα in length. Material. Holotype, CNU-ORT-LB2016067, female. Etymology. From the latin ‘ validus ’ for strong, refers to its unusually strong hind femur. Horizon and locality. Yixian Formation, Lower Cretaceous, Huangbanjigou Village, Beipiao City, Liaoning Province, China. General Description. A well-preserved female, the body medium size, head preserved much poorly and abdomen was been compressed. Head. Compound eye large, suborbicular. Thorax. Pronotum saddle-shaped with post margin prolongation. Prothorax leg preserved fragmentarily. Basal and middle part of metafemur much strong and distal part rapidly narrowed, inner side of it with irregular dark spots; hind knee robust, semi-lunar processes prominent on the dorsal edge; metatibia long and slender, dorsum with a row of tiny spines in basal half and two pairs of big leaf-like spurs in distal half visible; leaf-like spurs narrow, long and thickened, with a rachis in the middle part, then gradually thinned to the margin; basitarsomere slender, only preserved very basal part. Abdomen. Abdomen short, probably about half the length of the wing. Ovipositor gently recurved and elongated, its length more than two times than pronotum; the surface of the vavulae smooth. Wings. Forewing size large, Costa area relatively broad, CP reaching anterior margin about 1/3 length of forewing, with numerous branchlets; ScA reaching anterior margin at about origin of MP, with 3 branchlets connect with CP and distally simple; area between ScA and CP narrower than that of anterior and CP; ScP ending in anterior margin at about 1/2 length of forewing, several short branches connecting with ScA and four branches distally reaching anterior margin; stem R long and distinctly strong, branched into RA and RP near the ending of ScP; area between R and ScP relatively broad; RA ending in anterior margin close to apex and producing numerous oblique branches; RP with 11 comb-like branches, most of them reaching posterior margin, several distal terminals reaching anterior margin; most cross-veins between the branches of RP simple and straight; area between RA and RP narrow; M forking into MA and MP at about terminals of ScA; MA branching into MA1 and MA2 close to the last branch point of ScP; MA1 with 2 branches, with the first fused with RP and then forked immediately; MA2 simple; MP simple, originating at about terminal of ScA and reaching posterior margin at about 3/4 distal of forewing base; CuA+CuPaα simple, starting before the end point of CP, ending in posterior margin at about 2/3 length of forewing; CuPa straight, slanted toward the anterior margin, nearly as long as CuPaα; CuPaβ, CuPb and AA1 fused with each other at about 1/3rd of wing length; area between CuPb and AA1 relatively narrow, AA1 straight and strong; AA2 straight; area between AA1 and posterior margin filled with simple and straight cross-veins. Area between ScP and R, cross-veins between RA and RP, and that of CuPaβ, CuPb and AA1 are colored, one round dark spots located at the branching point of CuPa. Measurement. The specimen preserved very well but missing most part of head. The body total estimated length approximately 11 mm (from head to the end of abdomen without cercus and ovipositor). Pronotum 3.2 mm long. Left profemur preserved 2.2 mm long, protibia preserved 1.9 mm long. Left metathorax leg: metafemur preserved 5.6 mm long, metatibia 9.9 mm long and 0.5 mm wide, ds2 preserved 1.5 mm long and 0.4 mm wide, ds3 2.1 mm long and 0.4 mm wide, basitarsomere preserved 1.4 mm long. Right metathorax leg: metafemur preserved 7.7 mm long, metatibia 10.2 mm long and 0.5 mm wide, ds1 1.8 mm long, ds2 preserved 2.3 mm long and 0.4 mm wide. Ovipositor preserved 7.6 mm long. Left forewing 20.5 mm long and 4.5 mm wide (the maximum width). Remarks. This new species can be assigned to Elcaninae by its relatively narrow area between RA and RP, and CuPaβ, CuPb, and AA1 fused with each other. It falls in Panorpidium Westwood, 1854 by few branched ScA; three longitudinal branches between basal part of RP and CuA+CuPaα; and CuPaβ, CuPb and AA1 fused with each other distally (Fig. 3D–E). This new species is distinguished from all species of Panorpidium in the end of ScA close to the end of CP, CuPa straight and subequal to CuPaα in length and different pattern of coloration.

Panorpidium acusicaudum sp. nov. (Fig. 2) urn:lsid:zoobank.org:act: 7256D24D-2CFF-4080-A42F-45E3797C8BDC Diagnosis. Forewing medium size, CP short, reaching anterior margin basal of the fusion point of CuPaα and M+CuA, ScA and ScP with numerous branches, R forking into RA and RP after ScP reaching anterior margin, MP originating after ScA reaching anterior margin, CuPa much longer than CuPaα. Material. Holotype, CNU-ORT-LB2011041P/C, female. Etymology. From the latin ‘ acus ’ for needle and ‘ cauda ’ for tail, refers to its needle-like ovipositor. Horizon and locality. Yixian Formation, Lower Cretaceous, Huangbanjigou Village, Beipiao City, Liaoning Province, China. Description. A well-preserved female, the body medium size, head poorly preserved and abdomen vertically compressed. Head. Hypognathous, antennae filiform and slender, with more than 150 segments, flagellum much longer, flagellomere nearly rectangular, labrum approximates trapezoid. Thorax. Pronotum saddle-shaped, post margin not prolongated. Prothorax and mesothorax legs long and slender, pro- and mesotarsus both with 3 tarsomeres, basitarsomere longest, second tarsomere very small, distalmost tarsomere long and equal with basitarsomere. Metafemur much strong, upper side and outer surface of it with irregular dark spots; hind knee robust; metatibia long and slender, dorsum with three leaf-like spurs in distal half; distal of metatibia saved an apical spur; metatarsus with 3 tarsomeres, basitarsomere longest, ventral with two rows of tiny spines, second tarsomere very small, distalmost tarsomere long but about 1/2 length of the basitarsomere, claws well-developed. Abdomen. Abdomen short, about half the length of the wing. Ovipositor needle-like, elongated and sharply pointed, its length more than three times as long as pronotum; upper and lower vavulae almost of similar size, the surface of the vavulae smooth. Cerci conical, slender and unsegmented. Wings. Forewing large size, Costa area relatively narrow, CP short, reaching anterior margin before the fusion point of CuPaα and M+CuA, with numerous branchlets; ScA reaching anterior margin before origin of MP, with 6 branches distally reaching anterior margin; area between ScA and CP narrower than that of anterior and CP; ScP ending in anterior margin before R forking into RA and RP, several short branches basally connecting with ScA and numerous branches distally reaching anterior margin; stem R long and distinctly strong, branched into RA and RP near the mid-length of wing; area between R and ScP relatively broad; RA ending in anterior margin close to apex and producing numerous oblique branches; RP with 7 or 8 comb-like branches, most of them reaching posterior margin, several distal terminals reaching anterior margin; most cross-veins between the branches of RP simple and straight, some cross-veins between distal terminals dichotomic; area between RA and RP narrow; M forking into MA and MP between terminals of ScA and ScP; MA branching into MA1 and MA2 at the level of the end of ScP; MA1 with 2 branches, the first branch of it fused with RP; MP simple, reaching posterior margin at about 2/3 distal of forewing base; CuA+CuPaα simple, originating before the end of ScA and ending in posterior margin beyond mid-length of wing; CuPaβ, CuPb and AA1 fused with each other at about 1/3rd of wing length, basal of the ending of ScP; CuPa straight and long, more than three times as long as CuPaα, area between CuPb and AA1 relatively narrow, AA1 straight and strong; area between AA1 and posterior margin filled with irregularly reticulate veins. Area between RA and anterior margin, and that of CuPaβ, CuPb and AA1 are colored, one round dark spots located at the area of RP and MA1. Hindwing incomplete and strongly overlapped, basal part obscure; venation similar to that of forewing, RA with numerous branches ending in anterior margin; RP with 4–6 comb-like branches; MA and MP simple and long; CuA+CuPa straight, curved backward to the posterior at its terminal; other branches of CuP and anal veins are confounded because of fold and overlap, anals broad and with straight veins, covered by reticular transverse veins. Measurement. The estimated length of the body approximately 17 mm (from head to the end of abdomen). Left antennae preserved 55.2 mm long, right antennae preserved 18.3 mm long. Pronotum 2.6 mm long. Left Prothorax leg: profemur preserved 6.5 mm long and 0.9 mm wide, protibia preserved 5.5 mm long, basitarsomere 1.9 mm long, second tarsomere 0.3 mm long, distalmost tarsomere 1.6 mm long. Right Prothorax leg: profemur preserved 6.9 mm long and 0.9 mm wide, protibia preserved 6.4 mm long, basitarsomere 1.9 mm long, second tarsomere 0.3 mm long, distalmost tarsomere 1.7 mm long. Left mesothorax leg: mesotibia preserved 4.4 mm long, basitarsomere 1.7 mm long, second tarsomere 0.4 mm long, distalmost tarsomere 1.6 mm long. Right metathorax leg: metafemur 13.1 mm long, metatibia 11.6 mm long, ds2 preserved 3.0 mm long, apical spur 1.4 mm long, basitarsomere 3.3 mm long, distalmost tarsomere 1.4 mm long. Ovipositor 11.1 mm long. Left forewing 25.6 mm long and 5.0 mm wide (the maximum width), recognizable part of left hindwing preserved 18.6 mm long and 6.3 mm wide. Remarks. This new species can be assigned to Elcaninae by its relatively narrow area between RA and RP, and CuPaβ, CuPb, and AA1 fused with each other. It falls in Panorpidium Westwood, 1854 because of the following characters: few branched ScA; three longitudinal branches between basal part of RP and CuA+CuPaα; and CuPaβ, CuPb and AA1 fused with each other distally (Fig. 2C–D). The new species can be distinguished from P. parvum, P. bimaculatum and P. spica by the large size of body and wings, shorter CP with fewer branches, straight and longer CuPa and different pattern of coloration. It differs from P. tessellatum, P. proximum, P. sibiricum, P. yixianensis in CP with fewer branches, wider area between ScP and R, few branches of RP and different pattern of coloration., Published as part of Tian, He, Ren, Dong & Gu, Jun-Jie, 2022, Two new species of Elcaninae (Orthoptera, Elcanidae) recorded from the Early Cretaceous China, pp. 589-596 in Zootaxa 5128 (4) on pages 591-592, DOI: 10.11646/zootaxa.5128.4.8, http://zenodo.org/record/6480106

Genus Panorpidium Westwood, 1854 Type species. Panorpidium tessellatum Westwood, 1854 Included species. Panorpidium tessellatum Westwood, 1854; P. bimaculatum Gorochov, Jarzembowski and Coram, 2006; P. proximum Gorochov, Jarzembowski and Coram, 2006; P. sibiricum Sharov, 1968; P. yixianensis Fang, Wang, Zhang, Wang, Jarzembowski, Zheng, Zhang, Lia and Liu, 2015; P. spica Kim, Lee, Nam and Park, 2021; and possibly P. parvum Gorochov, Jarzembowski and Coram, 2006., Published as part of Tian, He, Ren, Dong & Gu, Jun-Jie, 2022, Two new species of Elcaninae (Orthoptera, Elcanidae) recorded from the Early Cretaceous China, pp. 589-596 in Zootaxa 5128 (4) on page 591, DOI: 10.11646/zootaxa.5128.4.8, http://zenodo.org/record/6480106, {"references":["Gorochov, A. V., Jarzembowski, E. A. & Coram, R. A. (2006) Grasshoppers and crickets (Insecta: Orthoptera) from the Lower Cretaceous of southern England. Cretaceous Research, 27, 641 - 662. https: // doi. org / 10.1016 / j. cretres. 2006.03.007","Sharov, A. G. (1968) Phylogeny of orthopteroid insects. Proceedings of the Paleontological Institute, Russian Academy of Sciences, 118, 1 - 217.","Kim, D. - Y., Lee, M., Nam, G. - S. & Park, T. - Y. S. (2021) The first orthopteran fossils from the Lower Cretaceous (Albian) Jinju Formation of Korea: Ethological implications for elcanids. Cretaceous Research, 125, 104843. https: // doi. org / 10.1016 / j. cretres. 2021.104843"]}

Panorpidium acusicaudum sp. nov. (Fig. 2) urn:lsid:zoobank.org:act: 7256D24D-2CFF-4080-A42F-45E3797C8BDC Diagnosis. Forewing medium size, CP short, reaching anterior margin basal of the fusion point of CuPaα and M+CuA, ScA and ScP with numerous branches, R forking into RA and RP after ScP reaching anterior margin, MP originating after ScA reaching anterior margin, CuPa much longer than CuPaα. Material. Holotype, CNU-ORT-LB2011041P/C, female. Etymology. From the latin ‘ acus ’ for needle and ‘ cauda ’ for tail, refers to its needle-like ovipositor. Horizon and locality. Yixian Formation, Lower Cretaceous, Huangbanjigou Village, Beipiao City, Liaoning Province, China. Description. A well-preserved female, the body medium size, head poorly preserved and abdomen vertically compressed. Head. Hypognathous, antennae filiform and slender, with more than 150 segments, flagellum much longer, flagellomere nearly rectangular, labrum approximates trapezoid. Thorax. Pronotum saddle-shaped, post margin not prolongated. Prothorax and mesothorax legs long and slender, pro- and mesotarsus both with 3 tarsomeres, basitarsomere longest, second tarsomere very small, distalmost tarsomere long and equal with basitarsomere. Metafemur much strong, upper side and outer surface of it with irregular dark spots; hind knee robust; metatibia long and slender, dorsum with three leaf-like spurs in distal half; distal of metatibia saved an apical spur; metatarsus with 3 tarsomeres, basitarsomere longest, ventral with two rows of tiny spines, second tarsomere very small, distalmost tarsomere long but about 1/2 length of the basitarsomere, claws well-developed. Abdomen. Abdomen short, about half the length of the wing. Ovipositor needle-like, elongated and sharply pointed, its length more than three times as long as pronotum; upper and lower vavulae almost of similar size, the surface of the vavulae smooth. Cerci conical, slender and unsegmented. Wings. Forewing large size, Costa area relatively narrow, CP short, reaching anterior margin before the fusion point of CuPaα and M+CuA, with numerous branchlets; ScA reaching anterior margin before origin of MP, with 6 branches distally reaching anterior margin; area between ScA and CP narrower than that of anterior and CP; ScP ending in anterior margin before R forking into RA and RP, several short branches basally connecting with ScA and numerous branches distally reaching anterior margin; stem R long and distinctly strong, branched into RA and RP near the mid-length of wing; area between R and ScP relatively broad; RA ending in anterior margin close to apex and producing numerous oblique branches; RP with 7 or 8 comb-like branches, most of them reaching posterior margin, several distal terminals reaching anterior margin; most cross-veins between the branches of RP simple and straight, some cross-veins between distal terminals dichotomic; area between RA and RP narrow; M forking into MA and MP between terminals of ScA and ScP; MA branching into MA1 and MA2 at the level of the end of ScP; MA1 with 2 branches, the first branch of it fused with RP; MP simple, reaching posterior margin at about 2/3 distal of forewing base; CuA+CuPaα simple, originating before the end of ScA and ending in posterior margin beyond mid-length of wing; CuPaβ, CuPb and AA1 fused with each other at about 1/3rd of wing length, basal of the ending of ScP; CuPa straight and long, more than three times as long as CuPaα, area between CuPb and AA1 relatively narrow, AA1 straight and strong; area between AA1 and posterior margin filled with irregularly reticulate veins. Area between RA and anterior margin, and that of CuPaβ, CuPb and AA1 are colored, one round dark spots located at the area of RP and MA1. Hindwing incomplete and strongly overlapped, basal part obscure; venation similar to that of forewing, RA with numerous branches ending in anterior margin; RP with 4–6 comb-like branches; MA and MP simple and long; CuA+CuPa straight, curved backward to the posterior at its terminal; other branches of CuP and anal veins are confounded because of fold and overlap, anals broad and with straight veins, covered by reticular transverse veins. Measurement. The estimated length of the body approximately 17 mm (from head to the end of abdomen). Left antennae preserved 55.2 mm long, right antennae preserved 18.3 mm long. Pronotum 2.6 mm long. Left Prothorax leg: profemur preserved 6.5 mm long and 0.9 mm wide, protibia preserved 5.5 mm long, basitarsomere 1.9 mm long, second tarsomere 0.3 mm long, distalmost tarsomere 1.6 mm long. Right Prothorax leg: profemur preserved 6.9 mm long and 0.9 mm wide, protibia preserved 6.4 mm long, basitarsomere 1.9 mm long, second tarsomere 0.3 mm long, distalmost tarsomere 1.7 mm long. Left mesothorax leg: mesotibia preserved 4.4 mm long, basitarsomere 1.7 mm long, second tarsomere 0.4 mm long, distalmost tarsomere 1.6 mm long. Right metathorax leg: metafemur 13.1 mm long, metatibia 11.6 mm long, ds2 preserved 3.0 mm long, apical spur 1.4 mm long, basitarsomere 3.3 mm long, distalmost tarsomere 1.4 mm long. Ovipositor 11.1 mm long. Left forewing 25.6 mm long and 5.0 mm wide (the maximum width), recognizable part of left hindwing preserved 18.6 mm long and 6.3 mm wide. Remarks. This new species can be assigned to Elcaninae by its relatively narrow area between RA and RP, and CuPaβ, CuPb, and AA1 fused with each other. It falls in Panorpidium Westwood, 1854 because of the following characters: few branched ScA; three longitudinal branches between basal part of RP and CuA+CuPaα; and CuPaβ, CuPb and AA1 fused with each other distally (Fig. 2C–D). The new species can be distinguished from P. parvum, P. bimaculatum and P. spica by the large size of body and wings, shorter CP with fewer branches, straight and longer CuPa and different pattern of coloration. It differs from P. tessellatum, P. proximum, P. sibiricum, P. yixianensis in CP with fewer branches, wider area between ScP and R, few branches of RP and different pattern of coloration.

Genus Lichenipolystoechotes Fang, Zheng & Wang, 2020 Type species. Lichenipolystoechotes angustimaculatus Fang, Zheng & Wang, 2020 Other included species. Lichenipolystoechotes ramimaculatus Fang, Ma & Wang, 2020, Published as part of Xu, Yifan, Zheng, Bingyu, Fang, Hui, Shih, Chungkun, Ren, Dong & Wang, Yongjie, 2021, New fossil giant lacewings from the Middle Jurassic Jiulongshan Formation of China (Neuroptera, Ithonidae), pp. 259-266 in Zootaxa 5067 (2) on page 262, DOI: 10.11646/zootaxa.5067.2.7, http://zenodo.org/record/5677973, {"references":["Fang, H., Labandeira, C. C., Ma, Y. M., Zheng, B. Y., Ren, D., Wei, X. L., Liu, J. X. & Wang, Y. J. (2020) Lichen mimesis in mid- Mesozoic lacewings. eLife, 9, e 59007. https: // doi. org / 10.7554 / eLife. 59007"]}

Lichenipolystoechotes fenestratus Xu, Fang, Shih & Wang, sp. nov. urn:lsid:zoobank.org:act: 65E67F61-A863-41F4-92EC-2BC07B1EFF91 (Figs. 2, 3) Etymology. The specific name fenestratus is a Latin word meaning “window-like”, referring to the diaphanous interior fenestrae in the forewing pattern of this species. Material. Holotype. CNU−NEU−NN2020003 (Fig. 2); Paratype. CNU−NEU−NN2020004 (Fig. 3). Type locality and horizon. Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Callovian-Oxfordian boundary, late Middle Jurassic. Diagnosis. Forewing broad, length-width ratio about 2.7; RP with about 36 branches; crossveins in radial sector rare except for two rows of outer and inner gradate series; RP1 deeply branched. Description. Forewing oval shaped, about 26 mm long, and 9.4 mm wide; trichosors well-defined; membrane with the distinctive disruptive markings, forming 14–15 transparent fenestrae along the margin and five large fenestrae in the middle; humeral veinlet recurrent; costal crossveins mostly simple except for the distal part; ScA present; ScP and RA fused distally; the region between ScP and RA relatively narrow, no crossveins present in between; eleven crossveins preserved between RA–RP area; RP with 36 close-set branches; crossveins in RP area relatively scarce, only one outer gradate series completely present, and the inner gradate series also partly present; the first branch of RP forked deeply with several bifurcated branches; M bifurcated basally, forming relatively abundant pectinate branches distally; CuA branched in middle, forming ca. 10 oblique pectinate branches; CuP simply biforked once, with distal bifurcated branches; A1 and A2 well-developed, forked three times each; A3 residually present. The paratype shows a lighter membrane with similar markings. The venations of holotype and paratype are basically the same with an exception of a plausible difference, i.e. the absence of gradate series of crossveins in paratype. However, this character is possibly not genuine because the crossveins in RP area are commonly difficultly visible owing to the overlapped wings. Remarks. The new species can be easily attributed to Lichenipolystoechotes for the typical markings and highly similar venation, i.e. the length of the wing, the presence of complicated disruptive markings and diverse fenestrae in forewing, the similar configurations of M branches, the multi-branched CuA with 7−11 branches and the bifurcated CuP (Fang et al., 2020). Nevertheless, the new species can be differentiated from the known Lichenipolystoechotes species for the number of RP branches (36 branches in the new species vs. 18 in L. angustimaculatus and 22 branches in L. ramimaculatus). In addition, the branched pattern of the first branch of RP in the new species is also different from other species with more complicated branches.

Fuscopolystoechotes reticulatus Xu, Zheng, Shih & Wang, sp. nov. urn:lsid:zoobank.org:act: EC1A65B9-8258-442B-896D-D62C415A225D (Fig. 1) Etymology. The specific name reticulatus is a Latin word meaning “reticulated”, referring to the reticulated pattern present on forewing of this species. Material. Holotype. CNU−NEU−NN2016005 (Fig. 1A–B); Paratype. CNU−NEU−NN2016008P/C (Fig. 1C), CNU−NEU−NN2016009 (Fig. 1D). Type locality and horizon. Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Callovian-Oxfordian boundary, late Middle Jurassic. Diagnosis. As for the genus due to monotypy. Description. Forewing elongated, ca. 39 mm long and 13 mm wide; membrane dark, with the distinct transparent patches along anterior and posterior margins; trichosors well-defined in most margin except for the proximal half of anterior margin; humeral vein recurrent, producing 4–5 short crossveins; costal crossveins with the distal forks, and absence of interlinked crossveins in between; space between Sc and RA relatively broad, sc-sa crossvein absent; ra-rp crossveins numerous; RP with 18 close-set branches, crossveins in RP area numerous and irregularly arranged, only one outer gradate series present; M forked close to the divergence of RP1 from RP; MA with 4 distal branches, MP with 6 distal branches; Cu proximally branched; CuA with about 9 oblique pectinate branches; CuP dichotomously branched; A1–A3 multi-dichotomously branched distally. Only forewing present, body features unknown.

Fuscopolystoechotes reticulatus Xu, Zheng, Shih & Wang, sp. nov. urn:lsid:zoobank.org:act: EC1A65B9-8258-442B-896D-D62C415A225D (Fig. 1) Etymology. The specific name reticulatus is a Latin word meaning ���reticulated���, referring to the reticulated pattern present on forewing of this species. Material. Holotype. CNU���NEU���NN2016005 (Fig. 1A���B); Paratype. CNU���NEU���NN2016008P/C (Fig. 1C), CNU���NEU���NN2016009 (Fig. 1D). Type locality and horizon. Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Callovian-Oxfordian boundary, late Middle Jurassic. Diagnosis. As for the genus due to monotypy. Description. Forewing elongated, ca. 39 mm long and 13 mm wide; membrane dark, with the distinct transparent patches along anterior and posterior margins; trichosors well-defined in most margin except for the proximal half of anterior margin; humeral vein recurrent, producing 4���5 short crossveins; costal crossveins with the distal forks, and absence of interlinked crossveins in between; space between Sc and RA relatively broad, sc-sa crossvein absent; ra-rp crossveins numerous; RP with 18 close-set branches, crossveins in RP area numerous and irregularly arranged, only one outer gradate series present; M forked close to the divergence of RP1 from RP; MA with 4 distal branches, MP with 6 distal branches; Cu proximally branched; CuA with about 9 oblique pectinate branches; CuP dichotomously branched; A1���A3 multi-dichotomously branched distally. Only forewing present, body features unknown., Published as part of Xu, Yifan, Zheng, Bingyu, Fang, Hui, Shih, Chungkun, Ren, Dong & Wang, Yongjie, 2021, New fossil giant lacewings from the Middle Jurassic Jiulongshan Formation of China (Neuroptera, Ithonidae), pp. 259-266 in Zootaxa 5067 (2) on pages 261-262, DOI: 10.11646/zootaxa.5067.2.7, http://zenodo.org/record/5677973

Genus Fuscopolystoechotes Xu, Zheng, Shih & Wang, gen. nov. urn:lsid:zoobank.org:act: 504F861B-6DE0-4C99-B962-0E12E3C83164 Type and only species: Fuscopolystoechotes reticulatus Xu, Zheng, Shih & Wang, sp. nov. Diagnosis. Forewing elongated, slightly broad with length-width ratio about 3; membrane fuscous, with the distinct transparent patches along the anterior and posterior margins; costal space narrow at base and then expanded to the max in proximal 1/5 of forewing, and subsequently narrowed gradually, with a distinctly shrinking region in distal half; humeral veinlet recurrent; presence of ScA; costal crossveins mostly biforked distally; absence of scp-ra crossvein; RP with more than 10 branches, RP1 deeply branched at the proximal half; crossveins in radial sector irregularly arranged except for the outer gradate series; M forked near the separated point of RP1 from RP; MA and MP bearing the similarly distal pectinate branches; Cu forked before the separated point of RP; CuA distinctly longer than CuP, forming the pectinate branches in distal; CuP with 3–4 pectinate branches, and each forming the secondary sub-branches; A1 dichotomously branched distally, A2 forked close to the base, A3 single branch, with several distal minor sub-branches. Etymology. The new generic name is a combination of the Latin prefix fusc- and polystoechotes (the representative genus of the Polystoechotes genus-group), referring to dark brown markings of forewing. The gender is masculine. Remarks. The new genus is clearly assigned to the Polystoechotes genus-group of Ithonidae for the combination of following characters: 1) forewing distinctly elongated (cf. relatively broad in Ithone genus-group); 2) absence of interlinked veinlets in between costal crossveins (cf. commonly present in Rapisma genus-group and Principiala genus-group); 3) Sc and RA fused distally (cf. generally separated in Ithone genus-groups and Rapisma genus-groups); 4) crossveins in radial sector regularly arranged or relatively sparse (cf. abundant and commonly irregularly arranged in other groups). The new genus is characterized by its special markings of forewing that is distinctly different from other genera of Polystoechotes genus-group except for Lichenipolystoechotes. However, the forewing markings in the new genus are distinctly broader and more regular comparing to Lichenipolystoechotes. In addition, their venation are also quite different, e.g., the branched costal crossveins, and the more complicated configuration of CuP, to justify setting up a new genus.

Lichenipolystoechotes fenestratus Xu, Fang, Shih & Wang, sp. nov. urn:lsid:zoobank.org:act: 65E67F61-A863-41F4-92EC-2BC07B1EFF91 (Figs. 2, 3) Etymology. The specific name fenestratus is a Latin word meaning ���window-like���, referring to the diaphanous interior fenestrae in the forewing pattern of this species. Material. Holotype. CNU���NEU���NN2020003 (Fig. 2); Paratype. CNU���NEU���NN2020004 (Fig. 3). Type locality and horizon. Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Callovian-Oxfordian boundary, late Middle Jurassic. Diagnosis. Forewing broad, length-width ratio about 2.7; RP with about 36 branches; crossveins in radial sector rare except for two rows of outer and inner gradate series; RP1 deeply branched. Description. Forewing oval shaped, about 26 mm long, and 9.4 mm wide; trichosors well-defined; membrane with the distinctive disruptive markings, forming 14���15 transparent fenestrae along the margin and five large fenestrae in the middle; humeral veinlet recurrent; costal crossveins mostly simple except for the distal part; ScA present; ScP and RA fused distally; the region between ScP and RA relatively narrow, no crossveins present in between; eleven crossveins preserved between RA���RP area; RP with 36 close-set branches; crossveins in RP area relatively scarce, only one outer gradate series completely present, and the inner gradate series also partly present; the first branch of RP forked deeply with several bifurcated branches; M bifurcated basally, forming relatively abundant pectinate branches distally; CuA branched in middle, forming ca. 10 oblique pectinate branches; CuP simply biforked once, with distal bifurcated branches; A1 and A2 well-developed, forked three times each; A3 residually present. The paratype shows a lighter membrane with similar markings. The venations of holotype and paratype are basically the same with an exception of a plausible difference, i.e. the absence of gradate series of crossveins in paratype. However, this character is possibly not genuine because the crossveins in RP area are commonly difficultly visible owing to the overlapped wings. Remarks. The new species can be easily attributed to Lichenipolystoechotes for the typical markings and highly similar venation, i.e. the length of the wing, the presence of complicated disruptive markings and diverse fenestrae in forewing, the similar configurations of M branches, the multi-branched CuA with 7���11 branches and the bifurcated CuP (Fang et al., 2020). Nevertheless, the new species can be differentiated from the known Lichenipolystoechotes species for the number of RP branches (36 branches in the new species vs. 18 in L. angustimaculatus and 22 branches in L. ramimaculatus). In addition, the branched pattern of the first branch of RP in the new species is also different from other species with more complicated branches., Published as part of Xu, Yifan, Zheng, Bingyu, Fang, Hui, Shih, Chungkun, Ren, Dong & Wang, Yongjie, 2021, New fossil giant lacewings from the Middle Jurassic Jiulongshan Formation of China (Neuroptera, Ithonidae), pp. 259-266 in Zootaxa 5067 (2) on page 262, DOI: 10.11646/zootaxa.5067.2.7, http://zenodo.org/record/5677973, {"references":["Fang, H., Labandeira, C. C., Ma, Y. M., Zheng, B. Y., Ren, D., Wei, X. L., Liu, J. X. & Wang, Y. J. (2020) Lichen mimesis in mid- Mesozoic lacewings. eLife, 9, e 59007. https: // doi. org / 10.7554 / eLife. 59007"]}

GENUS HIATOMEGOPS GUO ET AL. gen. nov. Zoobank registration: u r n: l s i d: z o o b a n k. org:act: F0561F03-E69D-45E6-A7FB-4A4D3BFEC6EE Etymology: The genus name is the combination of hiatus, opening in Latin, after the broad tracheal spiracle behind the epigastric furrow, and Lagonomegops, type genus of the family, from Greek -μεγόψ, magnification. The name is masculine. Type species: Hiatomegops spinalis sp. nov. Diagnosis: Palpal tibia and tarsus with several dorsal trichobothria; leg metatarsus slightly longer than tarsus; broad tracheal spiracle situated on postgastric area behind epigastric furrow. Remarks: Finding a fossil spider species in which both males and females have been described is rare. Among 19 reported genera in Lagonomegopidae, only Archaelagonops Wunderlich, 2012 is known from both male and female specimens. The two female specimens, which were subsequently included in Archaelagonops (erected on a male specimen originally), lack sufficient evidence to belong in that genus (Wunderlich, 2012, 2015). In general, female spiders are a larger size than conspecific males. Four lagonomegopid species, Lineaburmops beigeli, Lineaburmops maculatus Wunderlich, 2015, Parviburmops brevipalpus Wunderlich, 2015 and Planimegops parvus Wunderlich, 2017, comprised only of male specimens, were described as having a similar or smaller size with CNU-ARA-MA2020002, an adult female. CNU-ARA-MA2020002 can be distinguished from them by: carapace without white patch (carapace with distinct white patches in Lineaburmops beigeli and Lineaburmops maculatus); carapace much longer than wide (carapace length equal with width in Parviburmops brevipalpus); cephalic region of carapace with a pair of anterolateral protrusions next to the inner side of PME (protrusions absent in Planimegops parvus). We erected a new genus Hiatomegops based on CNU-ARA-MA2020002. Hiatomegops can be distinguished from the other two lagonomegopid genera (Lagonoburmops and Picturmegops) comprised only of female specimens by: body length 2.89 (body length about 8.00 in Lagonoburmops); carapace piriform in outline, much longer than wide (carapace distinctly narrowed posteriorly, slightly wider than long in Picturmegops); carapace without a stripe (carapace with distinct stripes in Picturmegops); metatarsus of leg I slightly longer than tarsus (metatarsus of leg I about two times longer than tarsus in Lagonoburmops). In addition, the holotypes of Grandoculus chemahawinensis Penney, 2004,? Lagonomegops cor Pérez-de la Fuente et al., 2013, Soplaogonomegops unzuei Pérez-de la Fuente et al., 2013, Spinomegops aragonensis Pérez-de la Fuente et al., 2013 and Spinomegops arcanus Pérez-de la Fuente et al., 2013 were identified as ‘juvenile or adult female’ (Penney, 2004; Pérez-de la Fuente et al., 2013). Hiatomegops can be easily separated from these by: carapace piriform in outline, without neck (carapace heart-shaped when viewed from above in? Lagonomegops cor; cephalic region constricted in a neck, with narrowest point medially in Soplaogonomegops unzuei), fovea small oval, deeply depressed (fovea absent in Spinomegops aragonensis and Spinomegops arcanus), leg I not enlarged, metatarsus and tarsus without scopulae (leg I distinctly longer and more robust than the others, metatarsus and tarsus with scopulae hairs in Grandoculus chemahawinensis)., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on pages 404-406, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Wunderlich J. 2012. On the fossil spider (Araneae) fauna in Cretaceous ambers, with descriptions of new taxa from Myanmar (Burma) and Jordan, and on the relationships of the superfamily Leptonetoidea. Beitrage zur Araneologie 7: 157 - 232.","Wunderlich J. 2015. On the evolution and the classification of spiders, the Mesozoic spider faunas, and descriptions of new Cretaceous taxa mainly in amber from Myanmar (Burma) (Arachnida: Araneae). Beitrage zur Araneologie 9: 21 - 408.","Wunderlich J. 2017. New and rare fossil spiders (Araneae) in mid Cretaceous amber from Myanmar (Burma), including the description of new extinct families of the suborders Mesothelae and Opisthothelae, as well as notes on the taxonomy, the evolution and the biogeography of the Mesothelae. Beitrage zur Araneologie 10: 72 - 279.","Penney D. 2004. Cretaceous Canadian amber spider and the palpimanoidean nature of lagonomegopids. Acta Paleontologica Polonica 49: 579 - 584.","Perez-De La Fuente R, Saupe EE, Selden PA. 2013. New lagonomegopid spiders (Araneae: † Lagonomegopidae) from Early Cretaceous Spanish amber. Journal of Systematic Palaeontology 11: 531 - 553."]}

LINEABURMOPS BEIGELI WUNDERLICH, 2015 (FIGS 4, 5) Wu n d e r l i c h (2 0 1 5): 2 5 5 –2 5 6, f i g s 2 3 6, 2 3 7, photos 104, 105. Material: Two adult males, specimen nos CNU-ARA-MA2020003 and CNU-ARA-MA2020004. Locality and horizon: Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, Mid-Cretaceous. Taphonomic features: CNU-ARA-MA 2020003: the pedicel and abdomen are somewhat broken; metatarsus, tarsus of left leg I, II and patella of left leg IV are missing; there are a few bubbles and impurities around the spider. A thrips near the tibia of left leg III is present as a syninclusion. CNU-ARA-MA2020004: the tarsus of left leg IV is incomplete, tarsal claws are missing; and there are many impurities and cracks around the spider. Description: Body length 2.98–3.08, dark-coloured, with a subtriangular white patch on clypeus, a pair of reniform white patches on the thoracic region and a pair of white longitudinal stripes on the abdomen dorsally (Figs 4A, B, 5F). Carapace length 1.57–1.59, width 0.96–0.99 at widest point, covered with short, dense, feathery setae pressed flat against the cuticle. Cephalic region of carapace slightly raised, with a pair of anterolateral protrusions next to the inner side of PME. Neck present between cephalic and thoracic regions, slightly narrower than cephalic region. Fovea absent. Eight eyes present; PME enormous, 0.17 in diameter, situated on anterolateral corner of carapace; PLE small, placed at the mediolateral margin of cephalic region, separated 0.25 from the PME centre; AME and ALE contiguous, situated between the PME and the clypeal margin. Chelicera length 0.58 and width 0.29 at base, cheliceral insertion distinctly separated from mouthparts, stridulatory files absent. Fang length 0.19. Chelicera with about eight peg teeth on promargin (Fig. 4C); retromargin not visible. Labium subtriangular, slightly longer than wide, not fused to sternum. Endites elongated, converging but not meeting in midline; serrula as a single row of teeth. Sternum covered with setae, convex and without tubercles. Palpal podomere lengths: fe 0.51, pa 0.18, ti 0.27, ta 0.67. Palp hairy, feathery setae at least present on tarsus. Tibia with three dorsal trichobothria in a single row (Fig. 5A). Cymbium elongate, finger-like distally (Fig. 5A). Detailed structures of male palp, such as embolus and conductor, not recognizable. Legs long but no legs enlarged; metatarsus distinctly much longer than tarsus in anterior legs; metatarsus slightly longer than tarsus in posterior legs. Leg formula I> II> IV> III: leg I cx 0.31, tr 0.07, fe 1.47–1.58, pa 0.34, ti 1.44–1.60, mt 1.10–1.16, ta 0.66–0.75; leg II cx 0.37, tr 0.07, fe 1.49–1.53, pa 0.31–0.37, ti 1.43–1.45, mt 1.08–1.11, ta 0.66–0.71; leg III cx 0.24–0.28, tr 0.06, fe 1.14–1.17, pa 0.30–0.32, ti 1.02–1.14, mt 0.67–0.69, ta 0.50–0.57; leg IV cx 0.29– 0.31, tr 0.08, fe 1.46–1.51, pa 0.32–0.35, ti 1.28–1.42, mt 0.83–0.86, ta 0.64–0.67. Metatarsus and tarsus without scopulae. Distal preening comb composed of six to eight short macrosetae, present on metatarsus of posterior legs ventrally (Fig. 5C). Feathery setae at least present on femur, patella, tibia and metatarsus (Fig. 5B). Tibiae at least with five, metatarsi at least with six, tarsi at least with nine trichobothria (Fig. 5C, D). Three tarsal claws, paired claws with five to eight teeth, median claw hook-like. Serrate accessory claw setae near the median claw present at least on left leg III (Fig. 5E). Abdomen ovoid, length 1.27–1.34, width 0.65– 0.68, densely covered with short feathery setae. Spinnerets and anal tubercle encircled by a quadrate glabrous band. Four spinnerets visible. ALS with two segments, distal segment much smaller than basal segment; PLS about as large as ALS, details not visible (Fig. 4E, F). Remarks: These two specimens can be assigned to Lineaburmops beigeli, the type species of genus Lineaburmops, by the following characters: the shape and placement of white patches on carapace and abdomen; the length of the podomeres., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on page 408, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Wunderlich J. 2015. On the evolution and the classification of spiders, the Mesozoic spider faunas, and descriptions of new Cretaceous taxa mainly in amber from Myanmar (Burma) (Arachnida: Araneae). Beitrage zur Araneologie 9: 21 - 408."]}

GENUS SCOPOMEGOPS GUO ET AL. gen. nov. Zoobank registration: urn: lsid: zoobank. org:act: A8809BE3-D927-432B-9C04-0AC83C27391F Etymology: The genus name is the combination of scopa, broom in Latin, after the clustered clavate spicules on the palpal retrolateral tibial apophysis (RTA), and the Greek suffix -μεγόψ, magnification, from Lagonomegops, type genus of the family. The name is masculine. Type species: Scopomegops fax sp. nov. Diagnosis: Male palpal tibia with a small apophysis on the distal retrolateral surface, several clustered clavate spicules situated on the top of retrolateral apophysis; leg metatarsus distinctly much longer than tarsus. Remarks: CNU-ARA-MA2 0 2 0 0 0 1 is an adult male. It can be easily distinguished from other male lagonomegopids by the apophysis on the distal retrolateral surface of the male palpal tibia. It cannot be included in the two lagonomegopid genera (Lagonoburmops Wunderlich, 2012 and Picturmegops Wunderlich, 2015) comprised only of female specimens, because of: carapace piriform in outline, much longer than wide (carapace distinctly narrowed posteriorly, slightly wider than long in Picturmegops); leg setae not long and dense (legs densely covered with long setae in Lagonoburmops). Conspecific spiders have some morphological differences at different developmental stages (Foelix, 2011). It is difficult to judge whether an adult male and a juvenile are the same species, especially in fossils. Therefore, we have erected a new genus Scopomegops based on CNU-ARA-MA2020001. Besides Scopomegops, seven species belonging to five genera of Lagonomegopidae are known to have one or more apophyses on the male palpal tibia. In Albiburmops annulipes Wunderlich, 2017, the carapace length is equal to its width, whereas the carapace is distinctly much longer than wide in Scopomegops. In Archaelagonops propinquus Wunderlich, 2015 and Archaelagonops scorsum Wunderlich, 2015, the palpal tibia bears two apophyses, whereas there is only one in Scopomegops. In Cymbiolagonops cymbiocalcar Wunderlich, 2015, the palpal tibia bears a divided apophysis, whereas the RTA is undivided in Scopomegops. In? Parviburmops bigibber Wunderlich, 2017, the tibial apophysis is situated on the ventral surface, but is on the retrolateral surface in Scopomegops. In Paxillomegops longipes Wunderlich, 2015 and? Paxillomegops brevipes Wunderlich, 2015, the palpal tibia bears numerous short peg bristles, which are lacking in Scopomegops., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on page 402, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Wunderlich J. 2012. On the fossil spider (Araneae) fauna in Cretaceous ambers, with descriptions of new taxa from Myanmar (Burma) and Jordan, and on the relationships of the superfamily Leptonetoidea. Beitrage zur Araneologie 7: 157 - 232.","Wunderlich J. 2015. On the evolution and the classification of spiders, the Mesozoic spider faunas, and descriptions of new Cretaceous taxa mainly in amber from Myanmar (Burma) (Arachnida: Araneae). Beitrage zur Araneologie 9: 21 - 408.","Foelix RF. 2011. Biology of spiders, 3 rd edn. New York: Oxford University Press.","Wunderlich J. 2017. New and rare fossil spiders (Araneae) in mid Cretaceous amber from Myanmar (Burma), including the description of new extinct families of the suborders Mesothelae and Opisthothelae, as well as notes on the taxonomy, the evolution and the biogeography of the Mesothelae. Beitrage zur Araneologie 10: 72 - 279."]}

ODONTOMEGOPS TITAN GUO & SELDEN, 2020 (FIG. 6) Guo et al. (2020): 2–4, figs 1, 2. Material: Male, specimen no. CNU-ARA-MA2019001 (holotype). Locality and horizon: Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, Mid-Cretaceous. Description: See Guo et al. (2020). Epigastric plate with more than 30 epiandrous spigots scattered along median margin of epigastric furrow (Fig. 6C). Tracheal spiracle broad, situated on postgastric area behind epigastric furrow, immediately dividing internally into two tracheae (Fig. 6A, B). Remarks: After regrinding and repolishing the holotype of Odontomegops titan, the male genitalia and tracheal spiracle are now shown clearly. They differ from the single, broad opening in Hiatomegops spinalis (Fig. 3G, H), in that two small openings are situated on the postgastric area behind the epigastric furrow in Odontomegops titan (Fig. 6A, B). We interpret the abdomen of CNU-ARA-MA2019001 as being somewhat modified and broken, thus exposing the internal part of the tracheal spiracle. Hence, we interpret the posterior respiratory system of Odontomegops titan as a single, broad tracheal spiracle, which immediately divides internally into two smaller tracheae., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on pages 410-411, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Guo XB, Selden PA, Shih CK, Ren D. 2020. Two new lagonomegopid spiders (Arachnida: Araneae) from the Mid- Cretaceous of northern Myanmar, with comments on the superfamilial placement of Lagonomegopidae. Cretaceous Research 106: 104257."]}

GENUS ODONTOMEGOPS GUO & SELDEN, 2020 Type species: Odontomegops titan Guo & Selden, 2020., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on page 410, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Guo XB, Selden PA, Shih CK, Ren D. 2020. Two new lagonomegopid spiders (Arachnida: Araneae) from the Mid- Cretaceous of northern Myanmar, with comments on the superfamilial placement of Lagonomegopidae. Cretaceous Research 106: 104257."]}

Guo, Xiangbo, Selden, Paul A., Ren, Dong (2022): New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae). Zoological Journal of the Linnean Society 195: 399-416, DOI: 10.1093/zoolinnean/zlab027, URL: https://www.mendeley.com/catalogue/e9144677-a36a-374f-9b9f-ef35c5f8cd3c/

GENUS SCOPOMEGOPS GUO ET AL. gen. nov. Zoobank registration: urn: lsid: zoobank. org:act: A8809BE3-D927-432B-9C04-0AC83C27391F Etymology: The genus name is the combination of scopa, broom in Latin, after the clustered clavate spicules on the palpal retrolateral tibial apophysis (RTA), and the Greek suffix -μεγόψ, magnification, from Lagonomegops, type genus of the family. The name is masculine. Type species: Scopomegops fax sp. nov. Diagnosis: Male palpal tibia with a small apophysis on the distal retrolateral surface, several clustered clavate spicules situated on the top of retrolateral apophysis; leg metatarsus distinctly much longer than tarsus. Remarks: CNU-ARA-MA2 0 2 0 0 0 1 is an adult male. It can be easily distinguished from other male lagonomegopids by the apophysis on the distal retrolateral surface of the male palpal tibia. It cannot be included in the two lagonomegopid genera (Lagonoburmops Wunderlich, 2012 and Picturmegops Wunderlich, 2015) comprised only of female specimens, because of: carapace piriform in outline, much longer than wide (carapace distinctly narrowed posteriorly, slightly wider than long in Picturmegops); leg setae not long and dense (legs densely covered with long setae in Lagonoburmops). Conspecific spiders have some morphological differences at different developmental stages (Foelix, 2011). It is difficult to judge whether an adult male and a juvenile are the same species, especially in fossils. Therefore, we have erected a new genus Scopomegops based on CNU-ARA-MA2020001. Besides Scopomegops, seven species belonging to five genera of Lagonomegopidae are known to have one or more apophyses on the male palpal tibia. In Albiburmops annulipes Wunderlich, 2017, the carapace length is equal to its width, whereas the carapace is distinctly much longer than wide in Scopomegops. In Archaelagonops propinquus Wunderlich, 2015 and Archaelagonops scorsum Wunderlich, 2015, the palpal tibia bears two apophyses, whereas there is only one in Scopomegops. In Cymbiolagonops cymbiocalcar Wunderlich, 2015, the palpal tibia bears a divided apophysis, whereas the RTA is undivided in Scopomegops. In? Parviburmops bigibber Wunderlich, 2017, the tibial apophysis is situated on the ventral surface, but is on the retrolateral surface in Scopomegops. In Paxillomegops longipes Wunderlich, 2015 and? Paxillomegops brevipes Wunderlich, 2015, the palpal tibia bears numerous short peg bristles, which are lacking in Scopomegops.

ODONTOMEGOPS TITAN GUO & SELDEN, 2020 (FIG. 6) Guo et al. (2020): 2–4, figs 1, 2. Material: Male, specimen no. CNU-ARA-MA2019001 (holotype). Locality and horizon: Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, Mid-Cretaceous. Description: See Guo et al. (2020). Epigastric plate with more than 30 epiandrous spigots scattered along median margin of epigastric furrow (Fig. 6C). Tracheal spiracle broad, situated on postgastric area behind epigastric furrow, immediately dividing internally into two tracheae (Fig. 6A, B). Remarks: After regrinding and repolishing the holotype of Odontomegops titan, the male genitalia and tracheal spiracle are now shown clearly. They differ from the single, broad opening in Hiatomegops spinalis (Fig. 3G, H), in that two small openings are situated on the postgastric area behind the epigastric furrow in Odontomegops titan (Fig. 6A, B). We interpret the abdomen of CNU-ARA-MA2019001 as being somewhat modified and broken, thus exposing the internal part of the tracheal spiracle. Hence, we interpret the posterior respiratory system of Odontomegops titan as a single, broad tracheal spiracle, which immediately divides internally into two smaller tracheae.

GENUS LINEABURMOPS WUNDERLICH, 2015 Type species: Lineaburmops beigeli Wunderlich, 2015., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on page 408, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Wunderlich J. 2015. On the evolution and the classification of spiders, the Mesozoic spider faunas, and descriptions of new Cretaceous taxa mainly in amber from Myanmar (Burma) (Arachnida: Araneae). Beitrage zur Araneologie 9: 21 - 408."]}

HIATOMEGOPS SPINALIS GUO ET AL. sp. nov. (FIG. 3) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 9168FFD9-29C1-4158-878B-D29E71594D41 Etymology: The specific name is the Latin adjective spinalis, meaning with spine(s), referring to the long macrosetae on the distal prolateral surface of palpal tibia. Holotype: Female, specimenno.CNU-ARA-MA2020002. Locality and horizon: Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, Mid-Cretaceous. Diagnosis: As for the genus. Ta p h o n o m i c f e a t u r e s: C N U - A R A - M A 2 0 2 0 0 0 2: completely preserved; many bubbles and impurities are present around the spider. Parts of insect legs are present as syninclusions. Description: Body length 2.89, carapace and legs darkcoloured, abdomen light-coloured, with an indistinct median dorsal stripe (Fig. 3A, B). Carapace piriform in outline, length 1.30, width 0.87 at widest point, covered with short, dense, feathery setae pressed flat against the cuticle (Fig. 3A). Cephalic region of carapace slightly raised, with a pair of anterolateral protrusions next to the inner side of PME (Fig. 3E). Fovea small oval, deeply depressed, situated in the centre of carapace, next to the posterior margin of cephalic region (Fig. 3A). Eight eyes present; PME enormous, 0.26 in diameter, situated on anterolateral corner of carapace; PLE small, placed at the mediolateral margin of cephalic region, separated 0.31 from the PME centre; AME and ALE contiguous, situated between the PME and the clypeal margin. Chelicera length 0.62, width 0.27 at base, cheliceral insertion close to mouthparts, stridulatory files absent. Fang length 0.22. Chelicera with about six peg teeth on promargin (Fig. 3E), retromargin not visible. Labium ligulate, slightly longer than wide, not fused to sternum. Endites elongated, converging and meeting in midline; serrula as a single row of teeth. Sternum shield shaped, covered with setae, convex and without tubercles. Palpal podomere lengths: fe 0.34, pa 0.17, ti 0.24, ta 0.45. Palp hairy; tibia and tarsus with several dorsal trichobothria; tibia with a long macrosetae on the distal prolateral surface (Fig. 3F). Palpal claw absent. Legs hairy, no legs enlarged; metatarsus slightly longer than tarsus. Leg formula II> I> IV> III: leg I cx 0.23, tr 0.07, fe 0.90, pa 0.47, ti 0.76, mt 0.64, ta 0.49; leg II cx 0.26, tr 0.07, fe 0.98, pa 0.40, ti 0.93, mt 0.63, ta 0.46; leg III cx 0.27, tr 0.07, fe 0.80, pa 0.36, ti 0.58, mt 0.47, ta 0.42; leg IV cx 0.33, tr 0.09, fe 0.93, pa 0.33, ti 0.73, mt 0.67, ta 0.47. Metatarsus and tarsus without scopulae. Distal preening comb composed of seven to ten short macrosetae, present on metatarsus of posterior legs ventrally. Feathery setae at least present on femur and patella of leg II. Tibiae with about ten, metatarsi with about six, tarsi with about six trichobothria in at least two rows. Three tarsal claws, paired claws with three to five teeth, median claw hook-like. Abdomen ovoid, length 1.33, width 0.89, densely covered with short, feathery setae. Female genitalia simple, epigyne absent. A broad subovoid opening, interpreted as tracheal spiracle, situated on the postgastric area behind the epigastric furrow (Fig.3G,H). Six spinnerets; ALS with three segments, basal segment much longer than others; PLS about as large as ALS, with two segments, distal segment almost as the equal length with basal segment; retrolateral surface of distal segment with a series of large spigots (Fig. 3I, J). Remarks: Lagonomegopidae was once thought to have a female-biased sex ratio (Pérez-de la Fuente et al., 2013), but as more and more male lagonomegopids are found and described, this issue was resolved (Park et al., 2019). The well-preserved female genitalia of CNU-ARA-MA2020002 shows that it is an unequivocal adult female., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on pages 406-408, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Perez-De La Fuente R, Saupe EE, Selden PA. 2013. New lagonomegopid spiders (Araneae: † Lagonomegopidae) from Early Cretaceous Spanish amber. Journal of Systematic Palaeontology 11: 531 - 553.","Park T-YS, Nam, K-S, Selden PA. 2019. A diverse new spider (Araneae) fauna from the Jinju Formation, Cretaceous (Albian) of Korea. Journal of Systematic Palaeontology 17: 1271 - 1297."]}

GENUS HIATOMEGOPS GUO ET AL. gen. nov. Zoobank registration: u r n: l s i d: z o o b a n k. org:act: F0561F03-E69D-45E6-A7FB-4A4D3BFEC6EE Etymology: The genus name is the combination of hiatus, opening in Latin, after the broad tracheal spiracle behind the epigastric furrow, and Lagonomegops, type genus of the family, from Greek -μεγόψ, magnification. The name is masculine. Type species: Hiatomegops spinalis sp. nov. Diagnosis: Palpal tibia and tarsus with several dorsal trichobothria; leg metatarsus slightly longer than tarsus; broad tracheal spiracle situated on postgastric area behind epigastric furrow. Remarks: Finding a fossil spider species in which both males and females have been described is rare. Among 19 reported genera in Lagonomegopidae, only Archaelagonops Wunderlich, 2012 is known from both male and female specimens. The two female specimens, which were subsequently included in Archaelagonops (erected on a male specimen originally), lack sufficient evidence to belong in that genus (Wunderlich, 2012, 2015). In general, female spiders are a larger size than conspecific males. Four lagonomegopid species, Lineaburmops beigeli, Lineaburmops maculatus Wunderlich, 2015, Parviburmops brevipalpus Wunderlich, 2015 and Planimegops parvus Wunderlich, 2017, comprised only of male specimens, were described as having a similar or smaller size with CNU-ARA-MA2020002, an adult female. CNU-ARA-MA2020002 can be distinguished from them by: carapace without white patch (carapace with distinct white patches in Lineaburmops beigeli and Lineaburmops maculatus); carapace much longer than wide (carapace length equal with width in Parviburmops brevipalpus); cephalic region of carapace with a pair of anterolateral protrusions next to the inner side of PME (protrusions absent in Planimegops parvus). We erected a new genus Hiatomegops based on CNU-ARA-MA2020002. Hiatomegops can be distinguished from the other two lagonomegopid genera (Lagonoburmops and Picturmegops) comprised only of female specimens by: body length 2.89 (body length about 8.00 in Lagonoburmops); carapace piriform in outline, much longer than wide (carapace distinctly narrowed posteriorly, slightly wider than long in Picturmegops); carapace without a stripe (carapace with distinct stripes in Picturmegops); metatarsus of leg I slightly longer than tarsus (metatarsus of leg I about two times longer than tarsus in Lagonoburmops). In addition, the holotypes of Grandoculus chemahawinensis Penney, 2004,? Lagonomegops cor Pérez-de la Fuente et al., 2013, Soplaogonomegops unzuei Pérez-de la Fuente et al., 2013, Spinomegops aragonensis Pérez-de la Fuente et al., 2013 and Spinomegops arcanus Pérez-de la Fuente et al., 2013 were identified as ‘juvenile or adult female’ (Penney, 2004; Pérez-de la Fuente et al., 2013). Hiatomegops can be easily separated from these by: carapace piriform in outline, without neck (carapace heart-shaped when viewed from above in? Lagonomegops cor; cephalic region constricted in a neck, with narrowest point medially in Soplaogonomegops unzuei), fovea small oval, deeply depressed (fovea absent in Spinomegops aragonensis and Spinomegops arcanus), leg I not enlarged, metatarsus and tarsus without scopulae (leg I distinctly longer and more robust than the others, metatarsus and tarsus with scopulae hairs in Grandoculus chemahawinensis).

ORDER ARANEAE CLERCK, 1757 FAMILY LAGONOMEGOPIDAE ESKOV & WUNDERLICH, 1995, Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on page 402, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Eskov KY, Wunderlich J. 1995. On the spiders from Taimyr ambers, Siberia, with the description of a new family and with general notes on the spiders from the Cretaceous resins. Beitrage zur Araneologie 4: 95 - 107."]}

HIATOMEGOPS SPINALIS GUO ET AL. sp. nov. (FIG. 3) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 9168FFD9-29C1-4158-878B-D29E71594D41 Etymology: The specific name is the Latin adjective spinalis, meaning with spine(s), referring to the long macrosetae on the distal prolateral surface of palpal tibia. Holotype: Female, specimenno.CNU-ARA-MA2020002. Locality and horizon: Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, Mid-Cretaceous. Diagnosis: As for the genus. Ta p h o n o m i c f e a t u r e s: C N U - A R A - M A 2 0 2 0 0 0 2: completely preserved; many bubbles and impurities are present around the spider. Parts of insect legs are present as syninclusions. Description: Body length 2.89, carapace and legs darkcoloured, abdomen light-coloured, with an indistinct median dorsal stripe (Fig. 3A, B). Carapace piriform in outline, length 1.30, width 0.87 at widest point, covered with short, dense, feathery setae pressed flat against the cuticle (Fig. 3A). Cephalic region of carapace slightly raised, with a pair of anterolateral protrusions next to the inner side of PME (Fig. 3E). Fovea small oval, deeply depressed, situated in the centre of carapace, next to the posterior margin of cephalic region (Fig. 3A). Eight eyes present; PME enormous, 0.26 in diameter, situated on anterolateral corner of carapace; PLE small, placed at the mediolateral margin of cephalic region, separated 0.31 from the PME centre; AME and ALE contiguous, situated between the PME and the clypeal margin. Chelicera length 0.62, width 0.27 at base, cheliceral insertion close to mouthparts, stridulatory files absent. Fang length 0.22. Chelicera with about six peg teeth on promargin (Fig. 3E), retromargin not visible. Labium ligulate, slightly longer than wide, not fused to sternum. Endites elongated, converging and meeting in midline; serrula as a single row of teeth. Sternum shield shaped, covered with setae, convex and without tubercles. Palpal podomere lengths: fe 0.34, pa 0.17, ti 0.24, ta 0.45. Palp hairy; tibia and tarsus with several dorsal trichobothria; tibia with a long macrosetae on the distal prolateral surface (Fig. 3F). Palpal claw absent. Legs hairy, no legs enlarged; metatarsus slightly longer than tarsus. Leg formula II> I> IV> III: leg I cx 0.23, tr 0.07, fe 0.90, pa 0.47, ti 0.76, mt 0.64, ta 0.49; leg II cx 0.26, tr 0.07, fe 0.98, pa 0.40, ti 0.93, mt 0.63, ta 0.46; leg III cx 0.27, tr 0.07, fe 0.80, pa 0.36, ti 0.58, mt 0.47, ta 0.42; leg IV cx 0.33, tr 0.09, fe 0.93, pa 0.33, ti 0.73, mt 0.67, ta 0.47. Metatarsus and tarsus without scopulae. Distal preening comb composed of seven to ten short macrosetae, present on metatarsus of posterior legs ventrally. Feathery setae at least present on femur and patella of leg II. Tibiae with about ten, metatarsi with about six, tarsi with about six trichobothria in at least two rows. Three tarsal claws, paired claws with three to five teeth, median claw hook-like. Abdomen ovoid, length 1.33, width 0.89, densely covered with short, feathery setae. Female genitalia simple, epigyne absent. A broad subovoid opening, interpreted as tracheal spiracle, situated on the postgastric area behind the epigastric furrow (Fig.3G,H). Six spinnerets; ALS with three segments, basal segment much longer than others; PLS about as large as ALS, with two segments, distal segment almost as the equal length with basal segment; retrolateral surface of distal segment with a series of large spigots (Fig. 3I, J). Remarks: Lagonomegopidae was once thought to have a female-biased sex ratio (Pérez-de la Fuente et al., 2013), but as more and more male lagonomegopids are found and described, this issue was resolved (Park et al., 2019). The well-preserved female genitalia of CNU-ARA-MA2020002 shows that it is an unequivocal adult female.

SCOPOMEGOPS FAX GUO ET AL. sp. nov. (FIG. 2) Zoobank registration: u r n: l s i d: z o o b a n k. org:act: A897B322-B55F-4652-AF1C-FAD42209F78C Etymology: The specific name is the Latin noun fax, meaning torch, referring to the shape of the male palp. Holotype: Male, specimen no. CNU-ARA-MA2020001. Locality and horizon: Hukawng Valley, Kachin State, northern Myanmar; lowermost Cenomanian, Mid-Cretaceous. Diagnosis: As for the genus. Taphonomic features: CNU-ARA-MA2020001: femur, patella, tibia, metatarsus, tarsus of left leg II and metatarsus, tarsus of right leg I are missing; left leg I, caused by developmental malformation or autotomy, is much smaller than right leg I; the abdomen is compressed; there is a large bubble on the left side of the abdomen, and several small bubbles around the spider. A thrips near the left leg IV is present as a syninclusion. Description: Body length 4.97, carapace light-coloured medially, dark-coloured laterally, a pair of indistinct dark longitudinal stripes on abdomen dorsally. Carapace piriform in outline, length 2.44, width 1.54 at widest point, covered with short, dense, feathery setae pressed flat against the cuticle. Cephalic region of carapace slightly raised, with a pair of distinct protrusions beneath the inner side of PME (Fig. 2G). Fovea oval, shallow, situated in the centre of carapace, next to the posterior margin of cephalic region. Eight eyes present; PME enormous, 0.43 in diameter, situated on anterolateral corner of carapace; PLE small, placed at the mediolateral margin of cephalic region, separated 0.50 from the PME centre; AME and ALE contiguous, situated between the PME and the clypeal margin (Fig. 2G). Chelicera length 0.96, and width 0.36 at base, cheliceral insertion close to mouthparts, stridulatory files absent. Fang length 0.33. Chelicera with about six peg teeth on promargin (Fig. 2C); retromargin unrecognizable. Labium ligulate, longer than wide, not fused to sternum. Endites elongated, converging and meeting in midline; apex and prolateral margin of endites brushy with dense setae; serrula as a single row of teeth. Sternum shield shaped, covered with setae, slightly convex and without tubercles. Palpal podomere lengths: fe 1.01, pa 0.40, ti 0.43, ta 0.98. Palp hairy, feathery setae at least present on tibia. Tibia with at least three dorsal trichobothria in a single row; a small apophysis present on the distal retrolateral surface of tibia, several clustered clavate spicules situated on the top of retrolateral apophysis (Fig. 2D, H). Male palp torch-like, cymbium elongate; embolus curved, slightly embraced by the long sheet conductor; median apophysis horn-like; subtegulum round, with a distally curved apophysis (Fig. 2I–L; File S4). Legs long but no legs enlarged; metatarsus distinctly much longer than tarsus in all legs. Leg formula II> I> IV> III: leg I cx 0.64, tr 0.11, fe 2.81, pa 0.92, ti> 2.69; leg II cx 0.60, tr 0.11, fe 2.95, pa 0.84, ti 3.42, mt 2.42, ta 0.87; leg III cx 0.60, tr 0.10, fe 1.94, pa 0.65, ti 1.92, mt 1.13, ta 0.63; leg IV cx 0.65, tr 0.12, fe 2.51, pa 0.77, ti 2.28, mt 2.01, ta 0.82. Metatarsus and tarsus without scopulae. Distal preening comb composed of eight short macrosetae, present on metatarsus of posterior legs ventrally (Fig. 2E, F). Feathery setae at least present on all tibia. Tibiae with about 14, metatarsi with about ten, tarsi with about ten trichobothria (Fig. 2E, F). Three tarsal claws, paired claws with about four to five teeth, distal two teeth distinctly larger than others, median claw hook-like. Abdomen length 2.17, width> 0.66, densely covered with short setae. Four spinnerets visible, details unrecognizable. Remarks: Holotype with left leg I much smaller than right leg I. It was caused by developmental malformation or autotomy. Autotomy is a voluntary act allowing most spiders to amputate one of their own legs to escape from a perilous situation. Although the lost legs can be replaced by new ones regenerated after the next moult, these are usually smaller, and need further moults to grow into normal size (Foelix, 2011)., Published as part of Guo, Xiangbo, Selden, Paul A. & Ren, Dong, 2022, New specimens from Mid-Cretaceous Myanmar amber illuminate the phylogenetic placement of Lagonomegopidae (Arachnida: Araneae), pp. 399-416 in Zoological Journal of the Linnean Society 195 on pages 402-404, DOI: 10.1093/zoolinnean/zlab027, http://zenodo.org/record/6599779, {"references":["Foelix RF. 2011. Biology of spiders, 3 rd edn. New York: Oxford University Press."]}

Hybristodryinus viriosus Wang, Olmi and Gao, sp. nov. LSID: urn:lsid:zoobank.org:act:79A30209-7D6E-4BAE-BA8B-31F7F 457D301 Etymology. The species is named after the Latin adjective viriosus (meaning, ���strong���). Material. Holotype: female. No. CNU-HYM-MA2015109. Diagnosis. Female of Hybristodryinus with occipital carina complete; antenna very slender, more than six times as long as head, reaching about second segment of metasoma; first flagellomere 19 times as long as broad; maxillary palpus very slender, with palpomere 4 about 20 times as long as broad. Type locality and horiz on. The amber specimen was collected from Kachin (Hukawng Valley) of northern Myanmar, which was dated at 98.79 �� 0.62 Ma, earliest Cenomanian (Cruikshank and Ko, 2003; Shi et al., 2012). Description. Female (Figs. 1 �� 3). Fully winged; length 6.7 mm (Figs. 1A, 1B, 1C, 2A). Head black, except clypeus and mandible testaceous; antenna testaceous; mesosoma black; metasoma brown; legs testaceous. Antenna 10-segmented, clavate, very slender, apparently hairless; lengths of antennomeres in following proportions: 20:8:46:36:29:19:14:13:10:12; antenna more than six times as long as head (200:33), reaching about second segment of metasoma. ADO's not visible. Flagellomeres much longer than wide; antennomere 9 about five times as long as broad (10:2); antennomere 3 about 19 times as long as broad (46:2.4). Head slightly convex, with sculpture not distinct; clypeus with anterior margin rounded (Figs. 2C, 2D, 2E); mandible 4-dentate (progressively larger from dorsal to ventral tooth); frons slightly convex, not longitudinally excavated medially, with transverse fold situated half way, not reaching eyes laterally; occipital carina complete, bordered by furrow, not reaching eyes laterally; vertex without longitudinal keels connecting lateral ocelli to occipital carina; occiput deeply excavated; eye bulging, bare, much shorter than head (22:33); POL = 13; OL = 5 (but head partly crushed, so this measure is the average between right and left OL); OOL = 15; TL = 8; OPL = 4; greatest breadth of lateral ocelli much shorter than POL (4:13); frontal line absent. Maxillary palpus with six palpomeres, number of labial palpomeres not observable, palpal formula probably 6/?. Maxillary palpus very slender, with palpomere 4 about 20 times as long as broad (10:0.5). Pronotum with deep anterior transverse impression between anterior collar and disc; disc humped, trapezoidal, not isosceles triangle shaped (Figs. 2C, 2D), not deeply longitudinally excavated medially; posterior collar absent; anterior surface of pronotum without longitudinal striae; sculpture of disc indistinct; pronotum longer than head (43:33); pronotal tubercle reaching tegula. Epicnemium present. Mesoscutum smooth, slightly shorter than pronotum (38:43). Notauli almost reaching posterior margin of mesoscutum (Fig. 2D); minimum distance between notauli slightly shorter than POL (10:13). Mesoscutellum with sculpture indistinct, shorter than mesoscutum (17:38). Metanotum shorter than scutellum (9:17), with sculpture indistinct. Metapectal-propodeal complex shorter than mesoscutum (30:38), reticulate rugose, areolae very wide, disc with posterior corners strongly projected posteriorly (Figs. 1B, 2A); propodeal declivity shorter than disc of metapectal-propodeal complex (10:20), indistinct, because hidden behind disc. Fore leg ratio: 38 (coxa): 41 (trochanter): 60 (femur): 70 (tibia): 38 (tarsomere 1): 5 (tarsomere 2): 8 (tarsomere 3): 40 (tarsomere 4): 51 (tarsomere 5); enlarged claw shorter than protarsomere 5 (37:51) (Figs. 1A, 1C, 3A, 3B, 3C). Protrochanter very slender and long, with long and slender proximal stalk, broadened after half-way; protrochanter about 13 times as long as broad (41:3.2); protarsomere 3 produced into hook; rudimentary claw present, much shorter than enlarged claw (7:37); arolium much shorter than enlarged claw (6:37); enlarged claw with distal apex sharp, with two subapical teeth and one row of small lamellae (number of lamellae not distinct). Protarsomere 5 with two rows composed of many lamellae, one row formed by longer lamellae (number of lamellae indistinct); distal apex with group of lamellae (number of lamellae indistinct). Mid leg ratio (Figs. 1A, 1C, 2A): 28 (coxa): 30 (trochanter): 56 (femur): 56 (tibia): 40 (tarsomere 1): 17 (tarsomere 2): 11 (tarsomere 3): 7 (tarsomere 4): 10 (tarsomere 5). Mesotrochanter short and broad, less than three times as long as broad: 15:6. Hind leg ratio (Figs.1A, 1C, 2A): 44 (coxa): 13 (trochanter): 67 (femur): 50 (tibia): 46 (tarsomere 1): 24 (tarsomere 2): 16 (tarsomere 3): 9 (tarsomere 4): 16 (tarsomere 5). Forewing hyaline, not darkened, with the usual venation of Dryininae (Figs. 3D, E); pterostigma much longer than broad (40:6), metacarp (PostabR1) shorter than pterostigma (25:40); marginal cell 2R1 open; stigmal veins (2r-rs & Rs) with distal part (Rs) much longer than proximal part (2r-rs) (39:21). The spurious vein of secondary nature that runs anterior of the claval furrow between cu-a and end of the 3-Cu. Forewing with usual three basal cells (costal (C), radial (R) and first cubital (1Cu)) clearly enclosed by pigmented veins. Hind wing not distinct. Metasomal petiole very short, much shorter than rest of metasoma (7:145). Metasoma slightly longer than mesosoma (145:140). Tibial spur formula 1/1/2. Male. Unknown. Host. Unknown. Remarks. Because protrochanter is about 13 times as long as broad and the disc of pronotum is not excised medially, the new species is similar to Hybristodryinus mon Perkovsky et al. (2019). The main difference between the new species and H. mon is that POL is much longer than OPL in H. viriosus and about as long as OPL in H. mon. Based on the description of Hybristodryinus viriosus sp. nov., the keys to the females of Hybristodryinus from Myanmar amber published by Perkovsky et al. (2019), Tribull et al. (2020) and Olmi et al., 2021, must be modified by replacing couplet 7 as follows: 7. Protrochanter less than seven times as long as broad (greatest breadth) ¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨ H. cretacicus Perkovskyet al. (2019) - Protrochanter about eight-thirteen times as long as broad (greatest breadth) ¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨ 8 8. Disc of pronotum deeply longitudinally excised medially (Fig. 2A in Tribull et al., 2020) ¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨ ¨¨¨¨¨¨¨¨¨¨¨¨¨ H. moutesoe Tribull et al. (2020). - Disc of pronotum not excised longitudinally medially (Figs 1B, 2D) ¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨ 9. 9. Head with POL about as long as OPL ¨¨¨¨¨¨¨¨¨¨¨ ¨¨¨¨¨¨¨¨¨¨¨¨¨¨¨ H. mon Perkovskyet al. (2019). - Head with POL much longer than OPL ¨¨¨¨¨¨¨¨¨¨¨ ¨¨¨¨¨¨¨¨¨¨¨ H. viriosus Wang, Olmi and Gao, sp. nov., Published as part of Wang, Zhen, Olmi, Massimo, Shih, Chungkun, Ren, Dong & Gao, Taiping, 2021, New species of Hybristodryinus from mid-Cretaceous Myanmar amber (Hymenoptera, Dryinidae), pp. 1-6 in Cretaceous Research 125 on pages 2-4, DOI: 10.1016/j.cretres.2021.104861, http://zenodo.org/record/4812995, {"references":["Cruikshank, R. D., Ko, K., 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences 21, 441 ‾ 455. https: // doi. org / 10.1016 / S 1367 - 9120 (02) 00044 - 5.","Shi, G., Grimaldi, D. A., Harlow, G. E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q., Li, X., 2012. Age constraint on Burmese amber based on UePb dating of zircons. Cretaceous Research 37, 155 ‾ 163. https: // doi. org / 10.1016 / j. cretres. 2012.03.014.","Perkovsky, E. E., Olmi, M., Muller, P., Martynova, K. V., 2019. A review of the genus Hybristodryinus Engel, 2005 (Hymenoptera, Dryinidae) from mid-Cretaceous Burmese amber, with a discussion on its phylogenetic significance. Cretaceous Research 99, 169 ‾ 189. https: // doi. org / 10.1016 / j. cretres. 2019.01.023.","Tribull, C., Barden, P., Olmi, M., 2020. Hybristodryinus moutesoe (Hymenoptera, Dryinidae), a new species from mid-Cretaceous Kachin (Burmese) amber. Cretaceous Research 114, 104528. https: // doi. org / 10.1016 / j. cretres. 2020.104528.","Olmi, M., Chen, H. - Y., Shih, C. K., Muller, P., Capradossi, L., Ren, D., Perkovsky, E. E., Guglielmino, A., 2021. New species of Hybristodryinus Engel (Hymenoptera, Dryinidae) from mid-Cretaceous amber of northern Myanmar, with notes on their possible hosts. Journal of Hymenoptera Research 81, 43 ‾ 55. https: // doi. org / 10.3897 / jhr. 81.57792."]}

Genus Hybristodryinus Engel, 2005 Type species: Hybristodryinus resinicolus Engel (2005), by monotype and original designation., Published as part of Wang, Zhen, Olmi, Massimo, Shih, Chungkun, Ren, Dong & Gao, Taiping, 2021, New species of Hybristodryinus from mid-Cretaceous Myanmar amber (Hymenoptera, Dryinidae), pp. 1-6 in Cretaceous Research 125 on page 2, DOI: 10.1016/j.cretres.2021.104861, http://zenodo.org/record/4812995, {"references":["Engel, M. S., 2005. A dryinine wasp in Burmese amber (Hymenoptera: Dryinidae). Polskie Pismo Entomologiczne 74, 485 ‾ 494."]}

Wang, Zhen, Olmi, Massimo, Shih, Chungkun, Ren, Dong, Gao, Taiping (2021): New species of Hybristodryinus from mid-Cretaceous Myanmar amber (Hymenoptera, Dryinidae). Cretaceous Research 125, No. 104861: 1-6, DOI: 10.1016/j.cretres.2021.104861

Hsiao, Yun, Li, Yun, Ren, Dong, Pang, Hong (2021): Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae). Zoological Journal of the Linnean Society 193: 1271-1293, DOI: 10.1093/zoolinnean/zlaa184, URL: https://doi.org/10.1093/zoolinnean/zlaa184

��� PALAEOCANTHARIS PANNA SP. NOV. (FIGS 5���6) Zo obank regi stra ti on: urn: ls id: zoo bank. org:act: A9078EA3-6759-42E4-A667-D1E9A6E77C33. Type material: Holotype: Female; no. CNU- COL-MA-0019; earliest Cenomanian, Hukawng Valley, northern Myanmar; deposited in the Key Laboratory of Insect Evolution & Environmental Changes, Capital Normal University in Beijing, China. Diagnosis: As for the genus. Description: Female. Body 5.8 mm long, 1.3 mm wide, parallel-sided, densely covered with suberect pubescence. Body yellowish brown, with abdomen lighter in colour (Fig. 5). Head (Fig. 6A, B) 0.9 mm long, 1.1 mm wide. Vertex smooth, shallowly depressed. Eyes small, hemispherically prominent, with fine facets, interocular distance 1.8 times maximum eye diameter. Anterior margin of clypeus arcuate, frontoclypeal suture absent. Labrum membranous, partially exposed. Gular sutures close, nearly confluent. Mandibles slender, strongly curved, with a sharp tooth at two thirds of inner margin. Apical segments of maxillary palpi and labial palpi flattened, securiform. Antennae (Fig. 5A) serrate; scape slender, widened apically; pedicel short, cylindrical; antennomeres 3���6 triangular; relative lengths of antennomeres 3���6: 1.0:1.3:1.4:1.4. Pronotum (Fig. 6C) 1.2 mm long, 1.1 mm wide, circular-shape. Anterior and lateral margins arcuate, posterior margin almost straight; disc densely punctate, with pair of shallow longitudinal grooves at basal half. Scutellum parallelsided, straight at apex. Elytra (Figs 5A, 6D) 3.9 mm long, 1.3 mm wide, parallel-sided, densely pubescent, with distinctly elevated humeri; lateral margins of elytra slightly elevated, with complete epipleuron; each elytron with eight longitudinal costae; costal interspaces with row of rounded cells; hindwing well-developed, completely concealed beneath elytra. Abdomen (Fig. 5B) weakly sclerotized, with seven ventrites; first five segments broad, as wide as elytra; last two segments (Fig. 6E) narrower, extending beyond elytra, apical tergite bilobed, ventrite 7 abruptly narrower distally, emarginate at apex. Female genitalia (Fig. 6E) retracted within abdomen, with pair of palpiform gonostyli exposed. Legs (Figs 5, 6F) slender; profemora strongly dilated, meso- and metafemora elongate, profemur length 0.9 mm, mesofemur 1.2 mm, and metafemur 1.9 mm; tibiae slender, nearly straight, with paired tibial spurs, protibia length 0.9 mm, mesotibia 1.5 mm and metatibia 2.0 mm; tarsal formula 5-5-5; tarsomeres 1���3 simple, tarsomere 1 longer than two times length of tarsomere 2; tarsomere 4 bilobed; claws simple, with minute dent at base. Etymology: The specific epithet is derived from pa���� ā, ���wisdom��� in the Pali language and a Buddhist term described as the understanding of the true nature of phenomena. We chose this name because the discovery of this species provided an insight into the palaeontology of Cantharidae. It is a noun in apposition., Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on pages 1283-1284, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254

† HUKAWNGICHTHYURUS MAHA SP. NOV. (FIGS 7–8) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 3F289BBE-89FF-414D-965C-691C8E06EC 3F. Type material: Holotype: Male; no. YL-COL00001; earliest Cenomanian, Hukawng Valley, northern Myanmar; deposited in the Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, China. Diagnosis: It can be distinguished from its congener by the head with distinct setae (Fig. 8A, B) (without setae in H. kyawkhaingwini); thick scape and enlarged antennomere 3–6 (Fig. 8C) (scape thin and elongate and antennomere 3–5 stout, only slightly enlarged in H. kyawkhaingwini); apical tergite moderately developed, with obtuse apex (Fig. 8E) (strongly developed,with acute apex in H.kyawkhaingwini);tibial spurs present (Fig. 8G) (absent in H. kyawkhaingwini). Description: Male. Body 3.0 mm long, 1.0 mm wide, parallel-sided, densely and finely pubescent, with pubescence distinct in head. Body yellowish brown, with dark brown elytra (Fig. 7). Head (Fig. 8A, B) 0.5 mm long, 0.5 mm wide. Vertex finely wrinkled and with setae, slightly convex, slightly narrowed behind the eyes. Eyes large, hemispherically prominent, with fine facets, interocular distance 2.2 times maximum eye diameter. Anterior margin of clypeus somewhat concave, frontoclypeal suture absent. Labrum subquadrate, membranous, exposed. Gular sutures close, nearly confluent. Mandibles elongate, strongly curved at apical fourth, with a tooth on inflexed outer margin. Apical segment of maxillary palpi flattened, elongate, securiform, with apex straight-truncated; apical labial palpomere triangular. Antennae (Fig. 8C) filiform, with central antennomeres enlarged, 11-segmented, extending to half of elytral length; scape thick, widened apically; pedicel short, stout, moniliform; antennomere 3–6 stout, enlarged, subequal in length, longer than pedicel; antennomeres 7–11 filiform; relative lengths of antennomeres 3–11: 1.0:1.2:1.2:1.2:1.2:1.1:1.1:1.1:1.2. Pronotum (Fig. 8D) 0.4 mm long, 0.6 mm wide, transverse. Anterior margin slightly arcuate, posterior margin slightly concave, lateral margins narrowed apically in anterior half; disc irregular, with a large depression anteriorly. Scutellum narrowed apically, truncate at apex. Elytra (Figs 7A, 8D) 2.5 mm long, 0.8 mm wide, subparallelsided, somewhat narrowed before the middle, rounded at apex; elytra smooth; hindwing well-developed, exposed beyond elytra. Abdomen (Fig. 7B) weakly sclerotized, with eight ventrites; first six segments broad, as wide as elytra; last two segments (Fig. 8E, F) narrower, apical tergite (Fig. 8E) distinctly furcate, ventrite 7 (Fig. 8F) with apical margin straight, ventrite 8 tapered apically, forming narrow, elongate process. Aedeagus (Fig. 8F) partially exposed, with pair of slender ventral processes, curved outwards. Legs (Fig. 8G) short; profemur length 0.9 mm, mesofemur 0.9 mm, and metafemur 0.9 mm; tibiae slender, nearly straight, with pair of tibial spurs, protibia length 0.7 mm, mesotibia 0.7 mm, and metatibia 0.8 mm; tarsal formula 5-5-5; tarsomeres 1–3 simple, tarsomere 1 as long as combined length of tarsomere 2 and 3; tarsomere 4 bilobed; claws simple, with minute dent at base. Etymology: The specific epithet is derived from m ā h ā, ‘great or noble’ in Sanskrit, referring to the discovery of this species being a great contribution to the palaeontology of Cantharidae. It is a noun in apposition. Remarks: This species is closely related to H. kyawkhaingwini with which it shares the small body size, the head shape with the small rostrum, the apical maxillary palpomere with straight-truncated apex, the filiform antennae with the short, stout and moniliform pedicel and the stout central antennomeres, the transverse pronotum, with large depression on the disc anteriorly, the smooth and distinctly shortened elytra, with largely exposed caudal segments and hindwings and the furcate caudal segment. Considering that only four minute differences are identified between H. kyawkhaingwini and H. maha, we place it in the genus Hukawngichthyurus, rather than propose a new genus.

† CRETOCANTHARIS GEN. NOV. (FIGS 3–4) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 99722309-6744-4BE2-9C5C-F85BDDEF6C68. Type species: † Cretocantharis veda. Diagnosis: Body small (c. 2.5 mm) (Fig. 3); eyes large, hemispherically prominent (Fig. 4A, B); apical maxillary palpomere bilaterally symmetric, securiform; mandibles simple, without tooth on the inner margin; antennae filiform, pedicel and antennomere 3 subequal in length (Fig. 3A). Pronotum subquadrate, parallel-sided, with a shallow median groove (Fig. 4C). Elytra slightly shortened, disc deeply punctate and reticulate, with longitudinal costae and inconspicuous cells (Fig. 4D). Legs elongate, with lobed tarsomere 4 and simple claws (Figs 3, 4F). Etymology: The name is composed of the prefix Creto derived from the Cretaceous and the Greek word κάνθαρισ, a beetle, derived from the genus Cantharis Linnaeus, 1758, type of Cantharidae. Gender: feminine.

† HUKAWNGICHTHYURUS FANTI & ELLENBERGER 2018 Type species: † Hukawngichthyurus kyawkhaingwini Fanti & Ellenberger, 2018: 202. Emended diagnosis (after Fanti & Ellenberger, 2018): Body small-sized (c. 3.0 mm); rostrum small; apical maxillary palpomere with apex straightly-truncated; antennae filiform, with pedicel short, stout and moniliform, central antennomeres (3–6) stout, and the remaining antennomeres filiform, extending to elytral midlength. Pronotum transverse, with large depression on the disc anteriorly. Elytra distinctly shortened, with caudal segments and hindwings largely exposed; disc smooth. Abdomen with caudal segment distinctly furcate, fishtail-shaped.

��� HUKAWNGICHTHYURUS FANTI & ELLENBERGER 2018 Type species: ��� Hukawngichthyurus kyawkhaingwini Fanti & Ellenberger, 2018: 202. Emended diagnosis (after Fanti & Ellenberger, 2018): Body small-sized (c. 3.0 mm); rostrum small; apical maxillary palpomere with apex straightly-truncated; antennae filiform, with pedicel short, stout and moniliform, central antennomeres (3���6) stout, and the remaining antennomeres filiform, extending to elytral midlength. Pronotum transverse, with large depression on the disc anteriorly. Elytra distinctly shortened, with caudal segments and hindwings largely exposed; disc smooth. Abdomen with caudal segment distinctly furcate, fishtail-shaped., Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on page 1284, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254, {"references":["Fanti F, Ellenberger S. 2018. A new fossil genus of soldier beetles (Coleoptera: Cantharidae) from Mid-Cretaceous Burmese amber: a probable case of adaptive convergence. Cretaceous Research 92: 201 - 204."]}

Palaeocantharis gen. nov.; Sanaungulus Fanti et al., 2018. General characteristics: Body small (2.6���6.0 mm); head prognathous to slightly deflexed (Figs 4A, B, 6A, B, 8A, B); frontoclypeal suture absent; eyes large, hemispherically prominent; apical maxillary and labial palpomeres bilaterally symmetric, securiform; mandibles usually toothed (Figs 6A, 8A, B), sometimes simple (Fig. 4A, B); gular sutures close, nearly confluent (Supporting Information, Fig. S1A���C); antennae 11-segmented. Pronotum simple, not elaborately sculptured, without lateral glandular pores (Figs 4C, 6C, 8D); mesopreepisternum contiguous; metaventrite with midline long, complete or near so, and branched. Elytra entirely covering abdomen (Fig. 5) or shortened in various degree, with abdomen and hindwings partially exposed (Figs 3, 7); disc usually deeply punctate, sometimes forming coarse reticulation (Figs 4D, 6D), or sometimes smooth (Fig. 8D). Tibial spurs usually present, rarely absent. Abdomen with large lateral eversible glandular pores (Supporting Information, Fig. S1L); ventrite 7 of male with apical margin usually entire or concave (Figs 4E, 8F), sometimes deeply emarginate; ventrite 8 of male symmetrical, forming narrow, elongate process. Aedeagus symmetrical (Figs 4E, 8F). Ovipositor with paired well-developed palpiform gonostyli (Fig. 6E)., Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on page 1281, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254

��� CRETOCANTHARIS GEN. NOV. (FIGS 3���4) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 99722309-6744-4BE2-9C5C-F85BDDEF6C68. Type species: ��� Cretocantharis veda. Diagnosis: Body small (c. 2.5 mm) (Fig. 3); eyes large, hemispherically prominent (Fig. 4A, B); apical maxillary palpomere bilaterally symmetric, securiform; mandibles simple, without tooth on the inner margin; antennae filiform, pedicel and antennomere 3 subequal in length (Fig. 3A). Pronotum subquadrate, parallel-sided, with a shallow median groove (Fig. 4C). Elytra slightly shortened, disc deeply punctate and reticulate, with longitudinal costae and inconspicuous cells (Fig. 4D). Legs elongate, with lobed tarsomere 4 and simple claws (Figs 3, 4F). Etymology: The name is composed of the prefix Creto derived from the Cretaceous and the Greek word ����������������, a beetle, derived from the genus Cantharis Linnaeus, 1758, type of Cantharidae. Gender: feminine., Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on page 1281, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254

��� HUKAWNGICHTHYURUS MAHA SP. NOV. (FIGS 7���8) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 3F289BBE-89FF-414D-965C-691C8E06EC 3F. Type material: Holotype: Male; no. YL-COL00001; earliest Cenomanian, Hukawng Valley, northern Myanmar; deposited in the Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, China. Diagnosis: It can be distinguished from its congener by the head with distinct setae (Fig. 8A, B) (without setae in H. kyawkhaingwini); thick scape and enlarged antennomere 3���6 (Fig. 8C) (scape thin and elongate and antennomere 3���5 stout, only slightly enlarged in H. kyawkhaingwini); apical tergite moderately developed, with obtuse apex (Fig. 8E) (strongly developed,with acute apex in H.kyawkhaingwini);tibial spurs present (Fig. 8G) (absent in H. kyawkhaingwini). Description: Male. Body 3.0 mm long, 1.0 mm wide, parallel-sided, densely and finely pubescent, with pubescence distinct in head. Body yellowish brown, with dark brown elytra (Fig. 7). Head (Fig. 8A, B) 0.5 mm long, 0.5 mm wide. Vertex finely wrinkled and with setae, slightly convex, slightly narrowed behind the eyes. Eyes large, hemispherically prominent, with fine facets, interocular distance 2.2 times maximum eye diameter. Anterior margin of clypeus somewhat concave, frontoclypeal suture absent. Labrum subquadrate, membranous, exposed. Gular sutures close, nearly confluent. Mandibles elongate, strongly curved at apical fourth, with a tooth on inflexed outer margin. Apical segment of maxillary palpi flattened, elongate, securiform, with apex straight-truncated; apical labial palpomere triangular. Antennae (Fig. 8C) filiform, with central antennomeres enlarged, 11-segmented, extending to half of elytral length; scape thick, widened apically; pedicel short, stout, moniliform; antennomere 3���6 stout, enlarged, subequal in length, longer than pedicel; antennomeres 7���11 filiform; relative lengths of antennomeres 3���11: 1.0:1.2:1.2:1.2:1.2:1.1:1.1:1.1:1.2. Pronotum (Fig. 8D) 0.4 mm long, 0.6 mm wide, transverse. Anterior margin slightly arcuate, posterior margin slightly concave, lateral margins narrowed apically in anterior half; disc irregular, with a large depression anteriorly. Scutellum narrowed apically, truncate at apex. Elytra (Figs 7A, 8D) 2.5 mm long, 0.8 mm wide, subparallelsided, somewhat narrowed before the middle, rounded at apex; elytra smooth; hindwing well-developed, exposed beyond elytra. Abdomen (Fig. 7B) weakly sclerotized, with eight ventrites; first six segments broad, as wide as elytra; last two segments (Fig. 8E, F) narrower, apical tergite (Fig. 8E) distinctly furcate, ventrite 7 (Fig. 8F) with apical margin straight, ventrite 8 tapered apically, forming narrow, elongate process. Aedeagus (Fig. 8F) partially exposed, with pair of slender ventral processes, curved outwards. Legs (Fig. 8G) short; profemur length 0.9 mm, mesofemur 0.9 mm, and metafemur 0.9 mm; tibiae slender, nearly straight, with pair of tibial spurs, protibia length 0.7 mm, mesotibia 0.7 mm, and metatibia 0.8 mm; tarsal formula 5-5-5; tarsomeres 1���3 simple, tarsomere 1 as long as combined length of tarsomere 2 and 3; tarsomere 4 bilobed; claws simple, with minute dent at base. Etymology: The specific epithet is derived from m ā h ā, ���great or noble��� in Sanskrit, referring to the discovery of this species being a great contribution to the palaeontology of Cantharidae. It is a noun in apposition. Remarks: This species is closely related to H. kyawkhaingwini with which it shares the small body size, the head shape with the small rostrum, the apical maxillary palpomere with straight-truncated apex, the filiform antennae with the short, stout and moniliform pedicel and the stout central antennomeres, the transverse pronotum, with large depression on the disc anteriorly, the smooth and distinctly shortened elytra, with largely exposed caudal segments and hindwings and the furcate caudal segment. Considering that only four minute differences are identified between H. kyawkhaingwini and H. maha, we place it in the genus Hukawngichthyurus, rather than propose a new genus., Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on pages 1284-1285, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254

† PALAEOCANTHARIS PANNA SP. NOV. (FIGS 5–6) Zo obank regi stra ti on: urn: ls id: zoo bank. org:act: A9078EA3-6759-42E4-A667-D1E9A6E77C33. Type material: Holotype: Female; no. CNU- COL-MA-0019; earliest Cenomanian, Hukawng Valley, northern Myanmar; deposited in the Key Laboratory of Insect Evolution & Environmental Changes, Capital Normal University in Beijing, China. Diagnosis: As for the genus. Description: Female. Body 5.8 mm long, 1.3 mm wide, parallel-sided, densely covered with suberect pubescence. Body yellowish brown, with abdomen lighter in colour (Fig. 5). Head (Fig. 6A, B) 0.9 mm long, 1.1 mm wide. Vertex smooth, shallowly depressed. Eyes small, hemispherically prominent, with fine facets, interocular distance 1.8 times maximum eye diameter. Anterior margin of clypeus arcuate, frontoclypeal suture absent. Labrum membranous, partially exposed. Gular sutures close, nearly confluent. Mandibles slender, strongly curved, with a sharp tooth at two thirds of inner margin. Apical segments of maxillary palpi and labial palpi flattened, securiform. Antennae (Fig. 5A) serrate; scape slender, widened apically; pedicel short, cylindrical; antennomeres 3–6 triangular; relative lengths of antennomeres 3–6: 1.0:1.3:1.4:1.4. Pronotum (Fig. 6C) 1.2 mm long, 1.1 mm wide, circular-shape. Anterior and lateral margins arcuate, posterior margin almost straight; disc densely punctate, with pair of shallow longitudinal grooves at basal half. Scutellum parallelsided, straight at apex. Elytra (Figs 5A, 6D) 3.9 mm long, 1.3 mm wide, parallel-sided, densely pubescent, with distinctly elevated humeri; lateral margins of elytra slightly elevated, with complete epipleuron; each elytron with eight longitudinal costae; costal interspaces with row of rounded cells; hindwing well-developed, completely concealed beneath elytra. Abdomen (Fig. 5B) weakly sclerotized, with seven ventrites; first five segments broad, as wide as elytra; last two segments (Fig. 6E) narrower, extending beyond elytra, apical tergite bilobed, ventrite 7 abruptly narrower distally, emarginate at apex. Female genitalia (Fig. 6E) retracted within abdomen, with pair of palpiform gonostyli exposed. Legs (Figs 5, 6F) slender; profemora strongly dilated, meso- and metafemora elongate, profemur length 0.9 mm, mesofemur 1.2 mm, and metafemur 1.9 mm; tibiae slender, nearly straight, with paired tibial spurs, protibia length 0.9 mm, mesotibia 1.5 mm and metatibia 2.0 mm; tarsal formula 5-5-5; tarsomeres 1–3 simple, tarsomere 1 longer than two times length of tarsomere 2; tarsomere 4 bilobed; claws simple, with minute dent at base. Etymology: The specific epithet is derived from paññ ā, ‘wisdom’ in the Pali language and a Buddhist term described as the understanding of the true nature of phenomena. We chose this name because the discovery of this species provided an insight into the palaeontology of Cantharidae. It is a noun in apposition.

��� CRETOCANTHARIS VEDA SP. NOV. (FIGS 3���4) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: F66A3FAF-BC8E-4101-A63D-93BC3A93D112. Type material: Holotype: Male; no. CNU008029; earliest Cenomanian, Hukawng Valley, northern Myanmar; deposited in the Key Laboratory of Insect Evolution & Environmental Changes, Capital Normal University in Beijing, China. Diagnosis: As for the genus. Description: Male. Body 2.6 mm long, 0.5 mm wide, parallel-sided, densely pubescent. Body yellowish brown, with dark brown head and elytra (Fig. 3). Head (Fig. 4A, B) 0.4 mm long, 0.6 mm wide. Vertex smooth, flat. Eyes large, hemispherically prominent, with fine facets, interocular distance 0.6 times maximum eye diameter. Anterior margin of clypeus straight, frontoclypeal suture absent. Labrum subquadrate, membranous, exposed. Gular sutures close, nearly confluent. Mandibles elongate, strongly curved at apical fourth, with smooth margins. Apical segments of maxillary palpi and labial palpi flattened, securiform. Antennae (Fig. 3A) filiform, 11-segmented, extending to half of elytral length; scape slender, widened apically; pedicel and antennomere 3 cylindrical, subequal in length; antennomeres 4���11 slightly compressed; relative lengths of antennomeres 3���11: 1.0:2.1:2.0:2.2:2.2:2.3: 2.2:2.2:2.3. Pronotum (Fig. 4C) 0.6 mm long, 0.4 mm wide, subquadrate. Anterior and posterior margins arcuate, lateral margins subparallel; disc slightly convex, with a shallow longitudinal median groove. Scutellum parallel-sided, straight at apex. Elytra (Figs 3A, 4D) 1.9 mm long, 0.5 mm wide, parallelsided, narrower posteriorly; each elytron with three longitudinal costae, stouter at humeral portion; costal interspaces with inconspicuous cells; hindwing well-developed, partially exposed beyond elytra. Abdomen (Fig. 3B) weakly sclerotized, with eight ventrites; first six segments broad, as wide as elytra; last two segments (Fig. 4E) narrower, ventrite 7 with apical margin concave, ventrite 8 with apical margin moderately emarginate. Aedeagus (Fig. 4E) partially exposed, with pair of slender processes curved dorsally, process widened apically, with pointed apex. Legs (Figs 3, 4F) slender; profemur length 0.6 mm, mesofemur 0.6 mm, and metafemur 0.7 mm; tibiae slender, nearly straight, with pair of tibial spurs, protibia length 0.7 mm, mesotibia 0.7 mm, and metatibia 0.9 mm; tarsal formula 5-5-5; tarsomeres 1���3 simple, tarsomere 1 as long as combined length of tarsomere 2 and 3; tarsomere 4 bilobed; claws simple, with minute dent at base. Etymology: The specific epithet is derived from the V��da (���the knowledge���), a collection of poems composed in archaic Sanskrit by Indo-European-speaking peoples who lived in north-west India during the 2 nd millennium BCE. We chose this name to refer to its discovery bringing invaluable knowledge to the palaeontology of Cantharidae. It is a noun in apposition., Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on pages 1282-1283, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254

KEY TO THE GENERA OF BURMITE CANTHARINAE1. Disc of pronotum without large depression; elytra deeply punctate, sometimes forming rows of reticulate punctures; caudal segment simple, unmodified.............................................................................................. 2 ��� Disc of pronotum with large depression anteriorly; elytra smooth; caudal segment distinctly furcated, fishtail-shaped.................................................................................................................. Hukawngichthyurus2. Pronotum broad and laterally rounded, sides forming continuous curve with anterior edge, without prominent anterior angle................................................................................................................................. 3��� Pronotum subquadrate to trapezoidal or distinctly transverse, sides not forming continuous curve with anterior edge because of noticeable anterior angle........................................................................................ 5 3. Antennae pectinate........................................................................................................................ Burmomiles��� Antennae filiform or serrate............................................................................................................................ 4 4. Antennae filiform; head slightly narrowed basally; pronotum broad anteriorly, with sides subparallel; tarsomeres 1���3 slightly bilobed.................................................................................................. Elektrokleinia ��� Antennae serrate; head strongly narrowed basally; pronotum strongly narrowed anteriorly; tarsomeres 1���3 simple................................................................................................................................. Palaeocantharis 5. Lateral edges of pronotum serrate......................................................................................... Myamalycocerus��� Lateral edges of pronotum simple................................................................................................................... 66. Pronotum trapezoidal or distinctly transverse............................................................................................... 7 ��� Pronotum subquadrate.............................................................................................................. Cretocantharis 7. Head rounded behind eyes; disc of pronotum rugose........................................................... Ornatomalthinus��� Head strongly narrowed basally; disc of pronotum smooth....................................................... Sanaungulus, Published as part of Hsiao, Yun, Li, Yun, Ren, Dong & Pang, Hong, 2021, Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae), pp. 1271-1293 in Zoological Journal of the Linnean Society 193 on page 1282, DOI: 10.1093/zoolinnean/zlaa184, http://zenodo.org/record/5753254