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Genus Litholingia Ren, 2002 Type species. Litholingia rhora Ren, 2002 p. 57. Included species. Litholingia rhora Ren, 2002 (type species); other species: Litholingi a eumorpha Ren, 2002, Litholingia longa Khramov, 2012, Litholingia polychotoma Ren, 2002, Litholingia ptesa Shi et al., 2011.

Litholinga rhora Ren, 2002 Revised diagnosis. Forewing with RP with branches distally forked (no branch deeply forked), with at least eight posterior branches; PCu terminated on the posterior margin beyond the second dichotomy of RP; CuA forks after the fork of CuP; CuA and CuP forked beyond the second dichotomy of RP; coloration pattern with alternating broad light and smaller dark stripes. Material. Specimen NIGP 177501, housed in the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Science, China. Locality and horizon. Middle-Upper Jurassic deposit of Daohugou, Haifanggou Formation, China. Description. Mesothorax massive, ca. 3.60 mm wide (distance between forewing insertions), and 3.09 mm long; other body parts not clearly visible. Leg with numerous small setae. Forewing 54.2 mm long and 18.1 mm wide. Forewing elongate, narrow and pubescent, the main longitudinal veins and wing margin scattered with minute trichobothria. Costal area relatively narrow, slightly wider than subcostal space, with only a series of gradate veins producing two rows of cells, a few simple crossveins near wing base. Subcostal veinlets unforked. ScP running parallel with RA. Dense crossveins present between ScP and RA. RP arising close to the base of the wings, with at least eight branches (some indistinct distally). First branch of RP arising at 0.13 of wing length, no branch of RP deeply forked, second dichotomy of RP located at midlength from first and third dichotomy, anterior to CuA fork and nearly aligned with CuP fork, nearly all branches of RP forked distally. Stem of M short with M fork slightly distal to first RP dichotomy, MA forked far distal from MP fork, MA branches forked near wing margin, with six apical branches. MP forking just after third dichotomy of RP and far from CuA from, with numerous branches forked near wing margin. CuA field larger than MP field forking after second dichotomy of RP and after CuP fork, with numerous branches forked near wing margin. CuP field smaller than CuA field, with five apical branches, CuP finishing near distal part of basal wing third. PCu long and pectinate, terminating slightly distal second dichotomy of RP. Wing base with two or three rows of cells distinctly present between A1 and A2. A2 short, abruptly bent toward wing margin. Dense cross venation between veins over entire wing. Trichosors clearly present along wing margin with several of them between each vein along the margin of the wing. Coloration pattern visible with alternating light and dark stripes, light ones (5���6) broader than dark ones, several additional white spots visible on sides of black bands between veins. Hind wing with a vein configuration similar to forewing, except for costal field with only one row of cells. Identification key to Litholingia species 1. CuP heavily branched (with five branches reaching wing margin), wings coloration pattern with thin dark stripes and broad white stripes............... Litholingia rhora - CuP pectinate or branched, if branched with less than five branches reaching wing margin, wings coloration pattern with broad dark stripes and thin white stripes.............................2 2. CuP pectinate............................................ Litholingia longa - CuP with a variable number of branches but never pectinate.....................................................................................3 3. Fourth branch of RP deeply forked, three branches of CuA originating at its fork..................... Litholingia polychotoma - Fourth branch of RP forked far distad its origin, two branches of CuA originating at its fork..............................4 4. Third branch of RP deeply forked, CuP likely with less than four branches, RP with seven branches..................................................... Litholingia ptesa - Third branch of RP forked far distad its origin, CuP with four branches, RP with at least nine branches............................................ Litholingia eumorpha, Published as part of Jouault, Corentin, Legendre, Fr��d��ric, Perrichot, Vincent, Condamine, Fabien L. & Nel, Andr��, 2022, Redescription of Litholingia rhora Ren, 2002 (Neuroptera: Grammolingiidae) from the Middle Jurassic of Daohugou, pp. 20-26 in Palaeoentomology 5 (1) on pages 22-23, DOI: 10.11646/palaeoentomology.5.1.3, http://zenodo.org/record/6280619, {"references":["Ren, D. (2002) A new lacewing family (Neuroptera) from the Middle Jurassic of Inner Mongolia, China. Insect Science, 9, 53 - 67. https: // doi. org / 10.1111 / j. 1744 - 7917.2002. tb 00172. x"]}

GENUS LONGITERMES JOUAULT, ENGEL, LEGENDRE & NEL GEN. NOV. Z o o b a n k r e g i s t r a t i o n: h t t p: / / z o o b a n k. org/ urn:lsid:zoobank.org:act: AB77601A-A6D5-4CB9- 9236-DEC7FF2144B7 Type species: Longitermes pulcher Jouault, Engel, Legendre & Nel sp. nov. Etymology: The generic name is a combination of the Latin longus, long, referring to the elongate head of the new species, and termes, termite, commonly used to refer to Isoptera. Gender masculine. Diagnosis: Imago. Head elongate, with anterior corners rounded orthogonal and posterior corners broadly rounded; occiput and frons convex; epicranial ecdysial line visible; ocelli absent; postclypeus trapezoidal; anteclypeus small; left mandible apical tooth elongate, sharp, three marginal teeth, LM 1 -LM 2 interdental space small, v-shaped, LM1 small, LM2 elongate, forming broad cutting edge, LM2-LM3 separated by a notch, LM3 bent toward base of mandible; right mandible with long, sharp apical tooth, and at least one marginal tooth of similar shape and length; pronotum slightly wider than head, anterior margin concave, posterior margin broadly convex with a median concavity, sides broadly convex; tarsi pentamerous (fourth tarsomere cryptic); tibial spur formula 3-4-2; all tibiae with strong setae but no additional spines; arolium present; forewing with scale large, with humeral margin nearly straight, claval suture straight (PCu), basal suture faintly arched; Sc, RA, RP heavily sclerotized; R branched within scale; RP nearly encompassing wing apex, with main branch bending to wing apex, with five secondary branches; M and CuA weaker than other surrounding veins; CuA with three branches in scales; anal veins absent; hindwing with delineated scale., Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on page 610, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105

MAGNIFITERMES KRISHNAI JOUAULT, ENGEL & NEL SP. NOV. (FIGS 6–8) Z o o b a n k r e g i s t r a t i o n: h t t p: / / z o o b a n k. o r g / urn:lsid:zoobank.org:act: DCEAE642-54ED-4FEC- 8AC4-CA6C23C821D8 Holotype: Specimen identifier IGR.BU-053 (a complete alate specimen preserved in a drop-shaped amber piece measuring 18 × 29 × 5 mm), housed in the amber collection of the Geological Department and Museum (IGR), Rennes, France. Etymology: The specific epithet honours the late Dr Kumar Krishna (1928–2014), eminent termitologist and beloved friend and collaborator of MSE. Dr Krishna’s work also inspired CJ to study fossil termites. He is sorely missed. The name is treated as a noun in the genitive case. Locality and age: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; Upper Albian to Lower Cenomanian, ‘Mid’-Cretaceous. Diagnosis: As for the genus (vide supra). Description: Imago. Body c. 13.5 mm long (as preserved and measured from tip of labrum to abdomen apex). Head massive, c. 2.87 mm long (measured from posterior margin to tip of labrum) and about 1.5 mm wide excluding compound eyes (measured beyond eyes), slightly trapezoidal, prognathous, sides convex, postclypeus and anteclypeus combined length c. 0.32mm long, labrum lobe-shape, sides nearly parallel, slightly narrower basally, apical margin nearly straight, with few setae; mandibles triangular shaped, c. 0.46 mm high, covered by labrum; left mandible apical tooth sharp, two marginal teeth, LM 1 -LM 2 interdental space broad, v-shaped, LM2 elongate; right mandible not visible; glossa (gl) and paraglossa (gl), lacinial incisor with at least one long tooth; five maxillary palpomeres visible, combined length c. 0.9 mm; three labial palpomeres present, at least 0.60 mm long; compound eyes, c. 0.52 mm long, circular, situated laterally near head midlength, and separated from posterior head margin by more than their length; ocelli present; fontanelle absent; antennae moniliform with 28(?) antennomeres. Pronotum massive, c. 1.58 mm long, trapezoidal, wider than head, anterior margin concave surrounding head, posterior margin broadly convex, with dorsal surface flatly arched in profile, sides convex. Legs slender; profemur, c. 1.37 mm long, protibia c. 0.90 mm long, protarsus c. 0.54 mm long; protibia with three spurs f1, f2, f3, plus additional spines (see Diagnosis, supra); mesotibia c. 1.31 mm long, mesotarsus c. 0.57 mm long; mesotibia with four observable spurs m1, m2, m3, m4, additional spines (see Diagnosis, supra); metatibia c. 1.87 mm long, with four observable spurs h1, h2, h3, h4, plus additional spines (see Diagnosis, supra), mesotarsus c. 0.75 mm long; all spurs flattened with serration; all tarsi pentamerous; pretarsal arolium present. Forewing scale at least 1.50 mm long. Hindwing hidden under forewing. Abdomen at least 6.46 mm long (damaged during fossilization), with at least nine observable segments; abdominal segments apparently slightly wider near midlength. Cerci with five cercomeres, stylus preserved but covered by a white coat. Colour: Not preserved. Remarks: Magnifitermes krishnai differs from most other Cretaceous Isoptera in combining a series of plesiomorphic characters in the wings (e.g. fine reticulation between veins in the scale and anal lobe present), tarsi wholly pentamerous (plesiomorphic character), a tibial spur formula of 3-4-4 and an overall massive trapezoidalshaped pronotum. The wholly pentamerous tarsi and enriched tibial spur formula suggest an earlydiverging position among Isoptera, among groups like Archotermopsidae, Hodotermitidae, Hodotermopsidae, Mastotermitidae and their various extinct relatives. Additionally, its general habitus, and particularly its large pronotum and anal lobe, suggest that M. krishnai is related to Mastotermitidae. Recently, Mastotermitidae were split into two distinct subfamilies Mastotermitinae and Idanotermitinae (Jiang et al., 2021). Idanotermitinae are mainly characterized by the absence of ocelli on the head of imagoes.Due to the presence of ocelli, the new fossil is placed in the subfamily Mastotermitinae. It differs from the other Mastotermitinae, in particular Mastotermes, by a distinct Y-shaped ecdysial scar. Potential paraphyly of Mastotermitinae relative to Idanotermitinae is in need of clarification., Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on pages 618-620, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105, {"references":["Jiang RX, Zhang HR, Eldredge KT, Song XB, Li YY, Tihelka E, Huang DY, Wang S, Engel MS, Cai C. 2021. Further evidence of Cretaceous termitophily: description of new termite hosts of the trichopseniine Cretotrichopsenius (Coleoptera: Staphylinidae), with emendations to the classification of lower termites (Isoptera). Palaeoentomology 4: 374 - 389."]}

GENUS MAGNIFITERMES JOUAULT, ENGEL & NEL GEN. NOV. Zoobank registration: http://zoobank.org/urn:lsid: z o o b a n k. o r g:a c t: B D E A7C 1 1- 0 2 9 6- 4 7F 8- A7 3 5 - 817A4D58F844 Etymology: The generic name is a combination of the Latin adjective magnificus, magnificent, and the noun termes, meaning ‘maggot’ or ‘termite’, and commonly used as the root for many genera of Isoptera. Gender masculine. Type species: Magnifitermes krishnai Jouault, Engel & Nel sp. nov. Diagnosis: Imago. Head massive, square-shaped, dorsally flattened, with posterior corners broadly rounded; epicranial scar present; ocelli present, located just above compound eyes (not separated from eyes by a space greater than their length); postclypeus trapezoidal and flat; anteclypeus anteriorly convex; left mandible with an apical and two distinct marginal teeth, LM1-LM2 interdental space broad, v-shaped, posterior margin of LM1 slightly longer than anterior margin LM2; pronotum massive, trapezoidal, wider than head, anterior margin concave surrounding head, posterior margin broadly convex, with dorsal surface flatly arched in profile, sides convex; tarsi pentamerous, with fifth tarsomere extremely elongate; tibial spur formula 3-4-4; protibia with two additional spines (fa, fb); mesotibia with four(?) additional spines (ma, mb, mc, md); metatibia with at least three additional spines (ha, hb, hc); pretarsal arolium present; forewing scale large, overlapping hindwing scale, claval suture (PCu) broadly arched, basal suture broadly arched; Sc, RA, RP, M heavily sclerotized; Sc with two main branches; R branched within scale; RP running parallel to costal margin, with at least five branches joining costal margin; M slightly closer to RP than to CuA, with at least four secondary branches radiating to apical margin; CuA less sclerotized, with several branches joining lower margin. Hindwing: scale (basal) suture absent; anal lobe (ano-jugal) developed (folded), Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on page 617, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105

GENUS ANISOTERMES ZHAO ET AL., 2019 Included species: Anisotermes xiai Zhao, Eggleton & Ren 2019 (type species) and A. bourguignoni Jouault, Engel, Huang & Nel sp. nov. Emended diagnosis: Head approximately rounded; ocelli absent; fontanelle absent; Y-shaped ecdysial line absent; mandible not exceeding labrum, with la and ra conspicuously longer than LM1 and RM1; antenna moniliform, with about 25 antennomeres, terminal eight antennomeres tapered; compound eyes hemispheric, lying at head midlength; pronotum definitely saddle-shaped, slightly wider than head in dorsal view; lateral margin weakly upcurved; procoxal carina present; additional tibial spines present on all legs; tibial spur formula 3-4-4; tarsi wholly pentamerous; arolium variable (absent in A. xiai but present in A. bourguignoni); wings membranous, densely reticulate; forewing scale large, overlapping hindwing base, basal suture convex, veins Sc, RA, RP and M more heavily pigmented than CuA. Forewing: all major veins originate within scale; Sc simple or double; RP with five to six main branches, terminating on costal margin anterior to wing apex; radial field of moderate width, parallel to costal margin; M running about midway between RP and CuA, terminating on posterior margin; medial field relatively narrow; CuA branched, terminating about same distance with Rs to wing apex; PCu (claval suture) arched, meeting basal suture on posterior margin. Hindwing without basal suture, large anal lobe present; M first inferior branch fading away, terminating in center of hindwing; cerci short, trimerous or tetramerous (with additional indented rings); abdominal styli absent in female but present in male., Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on page 613, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105, {"references":["Zhao Z, Eggleton P, Yin X, Gao T, Shih C, Ren D. 2019. The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites. Systematic Entomology 44: 612 - 623."]}

MASTOTERMES MYANMARENSIS JOUAULT SP. NOV. (FIGS 9, 10) Z o o b a n k r e g i s t r a t i o n: h t t p: / / z o o b a n k. o r g / urn:lsid:zoobank.org:act: F5CF600D-2068-424E-B255- 9F92051F952B Material: Holotype, IGR.BU-054, preserved in an elongate, pale and ovoid piece of amber measuring 50 × 20 × 9 mm, housed in the amber collection of the Geological Department and Museum (IGR), Rennes, France. Locality and horizon: Hkamti site, Hkamti District, Sagaing Region, Myanmar; Early Albian (c. 110 Mya), Early Cretaceous. Etymology: The specific epithet refers to the country of origin of the amber piece, Myanmar, combining the country name with the Latin toponymic adjectival suffix, – ēnsis, meaning ‘from’. Diagnosis: Imago. Wings membranous, long and broad, densely reticulate, with ‘cross-veins’ present; veins Sc, RA, RP and M more heavily sclerotized than CuA, reticulate veins obviously pigmented among radial and medial fields; forewing Sc with two main branches; R with two branches; width of radial field moderate, RA simple, RP with five main branches; medial field encompassing wing apex, M main vein closely parallel to RP main vein, with three main branches, first branch of M arising near apical-third of wing; CuA with five main branches, apical-most branch of CuA terminating into posterior margin and in apical-third of forewing length. Hindwing with large anal lobe; costal space wide; Sc simple, terminating on costal margin at basal-third of hindwing length; RA long, simple and terminating on costal margin before apical-third of hindwing length; RP with two main branches; medial field encompassing wing apex; M separating from RP near wing base, relatively closely parallel to RP, with three main branches, secondary branches present; CuA uniformly branching, terminating just posterior to wing apex. Description: Imago. Wings membranous, reticulate veins present; forewing scale part missing, forewing preserved part at least 10.70 mm long and 4.62 mm wide, apex rounded, middle section of posterior slightly convex basally then convex; veins Sc, RA, RP and M more heavily sclerotized than CuA and A; forewing Sc short, bifurcate at suture, posterior branch with a faint subbranch at margin; RA with one branch; radial field width moderate, occupying about one-sixth area on average; RP with five branches, proximal branch fading apically before anterior wing margin; M closely parallel to RP, with three main branches; CuA with numerous posterior branches, apical-most branch of CuA terminating on posterior margin well proximal to apex, more proximal branches loosely pectinate, apicalmost of these branches terminating on margin near wing midlength. Hindwing with large anal lobe, scale not preserved, 14.10 mm long and 6.0 mm wide with anal lobe and 5.16 mm wide without anal lobe; Sc longer than that of forewing; RA long, simple and terminating on costal margin before apical-third of hindwing length; RP with three branches, first branch originating just before point of contact between Sc and wing margin; M with four branches, separating from RP away from wing base, then closely parallel to RP, apical branch of M with a secondary branch; CuA with numerous posterior branches, some dichotomous, terminating posterior to wing apex. Colour: Not preserved. Remarks: Compared with the known ‘Mid’-Cretaceous fossil and the extant mastotermitids, M. myanmarensis stands out in having broadly rounded wings, wider intervals between the longitudinal veins (shared with M. monostichus Zhao et al., 2019), and a broad space above Sc on the hindwing. Additionally, the space occupied by Sc + RA + RP is extremely wide near the basal suture. Mastotermes myanmarensis has a hindwing with vein A1 without branches, while the extant M. darwiniensis has a vein A1 with numerous posterior branches (similar to M. monostichus), and the anal veins in the anal lobe are all simple (vs. with some dichotomies in M. darwiniensis and M. monostichus). Mastotermes myanmarensis can be differentiated from M. monostichus by the forewing with RA simple, M with only a reduced number of branches and the first dichotomy far from the wing base, CuA with more dichotomous branches; hindwing with a broad space above Sc, and the anal field with more branches. Mastotermes myanmarensis also differs from the extant M. darwiniensis by its temporal range (‘Mid’-Cretaceous vs. extant)., Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on pages 620-621, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105, {"references":["Zhao Z, Eggleton P, Yin X, Gao T, Shih C, Ren D. 2019. The oldest known mastotermitids (Blattodea: Termitoidae) and phylogeny of basal termites. Systematic Entomology 44: 612 - 623."]}

ANISOTERMES BOURGUIGNONI JOUAULT, ENGEL, HUANG & NEL SP. NOV. (FIGS 3–5) Z o o b a n k r e g i s t r a t i o n: h t t p: / / z o o b a n k. org/ urn:lsid:zoobank.org:act: A406BDD6-D799-4DB8- 93D0-4C2E61C71F1A Holotype: Specimen identifier NIGP177751 (a complete dealate specimen preserved in a rounded amber piece), housed in the collection of the Nanjing Institute of Geology and Palaeontology Chinese Academy of Sciences (NIGPAS), Nanjing, China. Etymology: The specific epithet honours Dr Thomas Bourguignon for his work on termite evolution. It is to be treated as a noun in the genitive case. Locality and age: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; Late Albian to Early Cenomanian, ‘Mid’-Cretaceous. Diagnosis: Imago. Head rounded, with anterior corners slightly orthogonal and posterior corners rounded; left mandible with la longer than LM1, both sharp; right mandible with an apical and two marginal teeth, ra shorter than RM 1, RM 1 and RM 2 separated by a sharp angle; occiput and frons convex; postclypeus trapezoidal; anteclypeus small; ocelli absent; pronotum massive, trapezoidal, wider than head, anterior margin concave surrounding head, posterior margin broadly convex, with dorsal surface flatly arched in profile, sides convex; tarsi pentamerous; tibial spur formula 3-4-4; protibia with two additional spine (fa, fb); mesotibia with six additional spines (ma-mf); metatibia with at least three additional spines (ha-hc); arolium present; forewing with scale large, with humeral margin welldefined and slightly arched, claval suture strongly curved (PCu), basal suture convex; Sc, RA, RP and CuA heavily sclerotized; Sc branched within scale and going outside of scale; R branched within scale; M weaker than other surrounding veins; CuA with three branches in scales; PCu strongly curved; anal veins not discernible within scale. Description: Imago. Body c. 10 mm long (from tip of labrum to abdomen apex). Head robust, c. 2.20 mm long (measured from under the pronotum to the tip of the labrum) and c. 1.93 mm wide excluding compound eyes, square-shaped in dorsal view, hypognathous, sides slightly convex, anterolateral corner slightly orthogonal, posterolateral corners rounded, Y-shaped ecdysial scar absent; tentorial pit visible (ttp); postclypeus (pc) 0.32 mm long, trapezoidal; labrum c. 0.50 mm long, lobe-shape, narrower basally; mandibles massive, triangular shaped, at least 0.53 mm high; buccal pieces with glossa (gl), paraglossa (gl), stipe (st), mentum (mt), submentum (sm) visible but not clearly describable; five maxillary palpomeres, combined length as preserved c. 1.2 mm; four(?) labial palpomeres present, with three discernible palpomeres, combined length c. 0.75 mm; compound eyes, c. 0.45 mm long, circular, situated laterally near head midlength, and separated from posterior head margin by more than their length; fontanelle not visible and probably absent; antenna moniliform with at least 22 antennomeres. Pronotum c. 1.60 mm long marginal teeth (number corresponding); sm, submentum; st, stipe; ttp, tentorial pit. Scale bars 0.5 mm (A, C, D); 1 mm (B). and c. 2.73 mm wide, overlapping head dorsally, sides slightly upturned. Legs robust; procoxa c. 1.38 mm long, profemur c. 1.51mm long, protibia c. 1.41mm long, protarsus at least 0.76 mm long; protibia with three spurs f1, f2, f3 and additional spines; mesocoxa c. 1.13 mm long, mesofemur medially swollen c. 1.57 mm long, mesotibia c. 1.72 mm long, mesotarsus c. 0.64 mm long; mesotibia with four observable spurs m1, m2, m3, m4 and additional spines; metacoxa at least 1.10 mm long, metafemur at least 2.24mm long, metatibia at least 1.50 mm long, with three observable spurs h1, h2, h3, h4 and additional spines; all spurs flattened with serration; all tarsi pentamerous; pretarsal arolium present. Forewing scale c. 2.36 mm long and 1.41 mm wide (measured distally), sclerotized, especially in anal area, with sparse setation. Hindwing scale not visible but proximal part of hindwing clearly cut beyond basal suture. Abdomen at least.6.0mm long with at least11observable segments; abdominal segments widest at midlength. Cerci multimerous c. 0.52 mm long, five to seven cercomeres (sometimes fused so not clearly countable). Colour: Not preserved. Remarks: Anisotermes bourguignoni differs from most other Cretaceous ‘lower’ Isoptera in having a plesiomorphic character in scale configuration (e.g. fine reticulation between veins in the scale), tarsi pentamerous (plesiomorphic character), a tibial spur formula of 3-4-4 and an overall massive trapezoidal-shaped pronotum. The wholly pentamerous tarsi and enriched tibial spur formula tend to imply an early diverged position within Isoptera, among groups like Archotermopsidae, Hodotermitidae, Mastotermitidae and their various extinct relatives. Additionally, its general habitus, and particularly its large pronotum, suggests that A. bourguignoni is related to Mastotermitidae or to taxa belonging to the ‘ Meiatermes -grade’. Interestingly, the hindwings of A. bourguignoni were cut slightly distal from the hindwing basal scale suture, while in the forewing the cut follows the basal suture. The hindwing basal scale suture was probably not fragile enough to break alone after the mating swarm, as it normally happens in a majority of extant termites. This suggests that the suture of the hindwing is rudimentary (symplesiomorphy) as in Mastotermes darwiniensis (Tillyard, 1931), and that the new specimen could be related to Mastotermitidae. Another character suggesting placement of Anisotermes within Mastotermitidae is the mandibular shape. In fact, Anisotermes and Mastotermes exhibit a similar configuration of teeth on both mandibles, and length variations (RM1 conspicuously longer than ra vs. equal in length) are usually indicative of different genera from the same family. Additionally, this placement is strengthened by the diagnostic characters of the new species: ocelli absent, procoxal carina present, arolium present and the aforementioned scale characters. The absence of ocelli indicates affinities with Idanotermitinae, but the new fossil differs from the genus Idanotermes Engel, 2008 in at least its different tibial spur formula (i.e. 3-4-4 vs. 3-4-3). The genus Anisotermes is here tentatively considered as a member of the Idanotermitinae (Jiang et al., 2021: 379); we confirm this placement mainly based on the presence of the anal lobe (characteristics of Mastotermitidae within the Isoptera) and based on the absence of ocelli. The new species differs from A. xiai by, inter alia, the presence of the pretarsal arolium and more additional spines along the tibiae., Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on pages 613-617, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105, {"references":["Krishna K, Grimaldi DA, Krishna V, Engel MS. 2013. Treatise on the Isoptera of the world. Bulletin of the American Museum of Natural History 377: 19891 - 12433.","Tillyard RJ. 1931. The wing venation of the order Isoptera: introduction and the family Mastotermitidae. Proceedings of the Linnean Society of New South Wales 56: 371 - 390.","Engel MS. 2008. Two new termites in Baltic amber (Isoptera). Journal of the Kansas Entomological Society 81 (3): 194 - 203.","Jiang RX, Zhang HR, Eldredge KT, Song XB, Li YY, Tihelka E, Huang DY, Wang S, Engel MS, Cai C. 2021. Further evidence of Cretaceous termitophily: description of new termite hosts of the trichopseniine Cretotrichopsenius (Coleoptera: Staphylinidae), with emendations to the classification of lower termites (Isoptera). Palaeoentomology 4: 374 - 389."]}

Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe, Nel, André (2022): Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera). Zoological Journal of the Linnean Society 196: 608-629, DOI: 10.1093/zoolinnean/zlac064

LONGITERMES PULCHER JOUAULT, ENGEL, LEGENDRE, & NEL SP. NOV. (FIGS 1, 2) Z o o b a n k r e g i s t r a t i o n: h t t p: / / z o o b a n k. org/ urn:lsid:zoobank.org:act: 6327861A-3757-4810- BEED-9800D20146C2 Material: Holotype, IGR.BU-052, preserved in an ovoid piece of amber measuring 18 × 12 × 2 mm, housed in the amber collection of the Geological Department and Museum (IGR), Rennes, France. Locality and horizon: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; Upper Albian to Lower Cenomanian, ‘Mid’-Cretaceous. Etymology: The specific epithet is from the Latin adjective pulcher, beautiful. Diagnosis: As for the genus (vide supra). Description: Imago specimen. Body c. 7.34 mm long (as preserved and measured from tip of labrum to abdominal apex). Head elongate, c. 1.44 mm long (measured from posterior margin to tip of labrum) and c. 0.82 mm wide excluding compound eyes (measured beyond eyes), trapezoidal in dorsal view, prognathous, sides slightly convex, postclypeus c. 0.11 mm long, anteclypeus c. 0.11 mm long, labrum lobe-shape, sides nearly parallel, slightly narrower basally, apical margin nearly straight, with few setae; mandibles triangular shaped, c. 0.20 mm high, covered by labrum; left mandible apical tooth elongate, sharp; three marginal teeth, LM1 small, LM1-LM2 interdental space small, v-shaped, LM 2 elongate, forming broad cutting edge, LM2-LM3 separated by a notch, LM3 bend toward base of mandible (general aspect similar to left imago mandible of Hodotermopsis); right mandible with long, sharp apical tooth, and at least one marginal tooth of similar shape and length; buccal pieces: glossa with two visible segments, paraglossa with three segments, length from base to apex (in mm) 0.05, 0.06, 0.1; lacinial incisor with two teeth; five maxillary palpomeres visible, length from base to apex (in mm): 0.06, 0.05, 0.08, 0.09, 0.12; three labial palpomeres present, total length as preserved c. 0.21 mm; compound eyes, c. 0.37 mm long, ovoid, situated laterally near head midlength, and separated from posterior head margin by more than their length; ocelli absent; fontanelle absent; antennae moniliform with 14 antennomeres. Pronotum c. 0.54 mm long and c. 1.0 mm wide, nearly flat with anterior margin concave, anterior corners arched, sides broadly convex and posterior margin convex with a medial indentation, posterior corners broadly arched. Legs robust; profemur medially swollen, c. 0.63 mm long, protibia c. 0.37 mm long, protarsus c. 0.22 mm long; protibia with three spurs f1, f2, f3; mesofemur medially swollen c. 0.63 mm long, mesotibia c. 0.42 mm long, mesotarsus c. 0.22 mm long; mesotibia with four observable spurs m1, m2, m3, m4; metafemur c. 0.51 mm long, metatibia c. 0.52 mm long, with two observable spurs h1, h2, mesotarsus c. 0.25 mm long; all tarsi pentamerous, with arolium. Forewing scale c. 0.80 mm long and 0.54 mm wide (measured distally), RP with additional tertiary branches joining costal margin; median (M) weak, closer to cubitus (Cu) than to radial sector (RP), with five branches terminating at lower margin; CuA with six primary and secondary branches extending to lower margin; anal vein absent. Hindwing (folded or damage): humeral suture weak, barely visible; wing venation apparently similar to forewing except anal vein present. Abdomen at least 4.0 mm long (damaged during the fossilization process) with at least ten observable segments; abdominal segments apparently slightly wider near midlength. Cerci multi-merous c. 0.16 mm long, five(?) cercomeres (sometimes fused so not clearly countable). Colour: Not preserved. Remarks: Longitermes pulcher is unique among Cretaceous termites, with an unprecedented combination of diagnostic characters making it close to the family Archotermopsidae, as treated in Jiang et al. (2021), i.e. encompassing Archotermopsis, Hodotermopsis and Zootermopsis. Here we follow the new system of Wang et al. (2022: table 1), which recently resolved paraphyly of Archotermopsidae by elevating Hodotermopsinae to family rank. In fact, L. pulcher shares with Archotermopsidae and Hodotermopsidae the lack of a fontanelle; a Y-shaped ecdysial line present; large compound eyes; ocelli absent; pronotum faintly arched, slightly wider than head; left mandible with an apical tooth and three marginal teeth. Legs: five tarsomeres (some cryptic); tibial spur formula 3-4-2; arolium present. Cerci with five(?) cercomeres. Forewing: costal margin flat to faintly arched; humeral suture well defined, almost straight; Sc, RA, RP sclerotized, RP with the main branch bending to apex of wing, five to six secondary branches and additional tertiary branches joining costal margin; M weak, with secondary branches terminating at lower margin; Cu with primary and secondary branches extending to lower margin; anal vein absent. The wing venation with the radial field encompassing most of the wing apex and the shape of the left mandibular teeth, nearly similar to that of Hodotermopsis, strengthen the taxonomic placement. However, the phylogenetic position of the new genus is hard to ascertain and it may be a stem representative of Hodotermopsidae or of Archotermopsidae (sensu Wang et al., 2022). Since the new specimen differs from the genera Archotermopsis and Zootermopsis (Archotermopsidae sensu Wang et al., 2022), at least based on its mandibular shape with LM2 elongate and forming a broad cutting edge (vs. sharp and not forming a cutting edge in the aforementioned genera) (Krishna et al., 2013), and resembles more the genus Hodotermopsis in its wing venation and mandibular configuration, we presume that Longitermes is probably a stem-Hodotermopsidae (sensu Wang et al., 2022). Hodotermopsis differs from the new specimen in possessing small eyes, while they are large in Longitermes, but also in having numerous antennomeres (at least 23), while the new specimen only possesses 14 (the antenna might not be complete but the number of antennomeres seems to be lower than in Hodotermopsis) (Krishna et al., 2013). Recently, several families were erected based on Cretaceous specimens from Kachin amber (e.g. Melqartitermitidae and Mylacrotermitidae) but our specimen cannot be placed in any of them (Jiang et al., 2021). Its broad radial field precludes affinities with the Melqartitermitidae and the presence of a welldefined forewing basal suture demarcating the scale precludes affinities with Mylacrotermitidae (Jiang et al., 2021). Similarly, the new specimen differs from representatives of the family Krishnatermitidae, at least, because of its forewing with PCu straight (vs. arched in Krishnatermes), RP encompassing the wing apex (variable in Krishnatermes) and its hindwing with a defined scale (vs. absent in Krishnatermes). However, the habitus of Longitermes and Krishnatermes are at first sight similar, which, if not symplesiomorphic, may indicate a proximity between the two genera and a hypothetical proximity of Krishnatermes with the Teletisoptera., Published as part of Jouault, Corentin, Engel, Michael S., Fls, Legendre, Frédéric, Huang, Diying, Grandcolas, Philippe & Nel, André, 2022, Incrementing and clarifying the diversity and early evolution of termites (Blattodea: Isoptera), pp. 608-629 in Zoological Journal of the Linnean Society 196 on pages 610-612, DOI: 10.1093/zoolinnean/zlac064, http://zenodo.org/record/7195105, {"references":["Jiang RX, Zhang HR, Eldredge KT, Song XB, Li YY, Tihelka E, Huang DY, Wang S, Engel MS, Cai C. 2021. Further evidence of Cretaceous termitophily: description of new termite hosts of the trichopseniine Cretotrichopsenius (Coleoptera: Staphylinidae), with emendations to the classification of lower termites (Isoptera). Palaeoentomology 4: 374 - 389.","Wang M, Hellemans S, Sobotnik J, Arora J, Bucek A, Sillam-Dusses D, Clitheroe C, Lu T, Lo N, Engel MS, Roisin Y, Evans TA, Bourguignon T. 2022. Phylogeny, biogeography, and classification of Teletisoptera (Blattaria: Isoptera). Systematic Entomology. In Press. Doi: 10.1111 / syen. 12548","Krishna K, Grimaldi DA, Krishna V, Engel MS. 2013. Treatise on the Isoptera of the world. Bulletin of the American Museum of Natural History 377: 19891 - 12433."]}

GENUS KERATODELLITHA JOUAULT, MARÉCHAL, WANG & PERRICHOT GEN. NOV. 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: 1CFB1DFA-B7F1-4414-8CAD-67515F278530 Type species: Keratodellitha basilisci Jouault, Maréchal, Wang & Perrichot, sp. nov. Other species included: Keratodellitha anubis Jouault, Maréchal, Wang & Perrichot sp. nov.; Keratodellitha kirin Maréchal, Jouault & Perrichot sp. nov. Etymology: The new generic name is a Latinized combination of the Greek words Κ έρας, horn, and δελλήθως, a kind of wasp. The gender of the name is feminine. Diagnosis: Head with vertex flat to slightly convex; cephalic horn without apical teeth, with two transverse and parallel ridges, anterior ridge being slightly wider than second but both similar in shape; compound eye ovoid, without circumocular carina, inner lateral margins slightly concave; toruli widely separated by cephalic horn, located on lateral surfaces, opening dorsolaterally; malar space shorter than width of mandibular base; mandible with apical teeth slightly pointing, not overlapping tooth of opposite mandible; gena shorter than compound eyes; antenna with 22 flagellomeres, flagellum gradually decreasing in length and slightly in width towards apex; pronotum extending forward as short neck; legs bearing tarsal plantulae; fore wing with lcu-a confluent to 1M; veins M+Cu et Cu aligned; 2m-cu tubular; 2cu-a and 2A present; hind wing with numerous hamuli (five or more)., Published as part of Jouault, Corentin, Maréchal, Arthur, Condamine, Fabien L., Wang, Bo, Nel, André, Legendre, Frédéric & Perrichot, Vincent, 2022, Including fossils in phylogeny: a glimpse into the evolution of the superfamily Evanioidea (Hymenoptera: Apocrita) under tip-dating and the fossilized birth-death process, pp. 1396-1423 in Zoological Journal of the Linnean Society 194 on page 1402, DOI: 10.1093/zoolinnean/zlab034, http://zenodo.org/record/6458537

KERATODELLITHA BASILISCI JOUAULT, MARÉCHAL, WANG & PERRICHOT SP. NOV. (FIGS 1, 2) 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: C67E5240-F1BC-4836-86F7-3B9D1D375AA8 Holotype: NIGP174738, preserved in a rectangular piece of amber measuring 17 mm × 10 mm × 6 mm. Type locality and horizon: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; upper Albian to lower Cenomanian, mid-Cretaceous. Etymology: The specific epithet refers to the basilisk (Latin basiliscus), a mythical animal that intermingled the traits of several animals, and is a general allusion to the fierce and enigmatic habitus of the species. The specific epithet is a noun in a genitive case. Diagnosis: Scape long, ~3.5 times longer than pedicel; outer (external) pretarsal claw of foreleg with two preapical teeth; fore wing vein Rs+M as long as 2Rs; third submarginal cell as long as second one; 1rs-m meeting Rs slightly before r-rs; 2rs-m tubular; second discal cell short (twice as long as wide); hind wing with six hamuli; gaster laterally compressed (as in Evaniidae), rounded in shape. Description: Female. Total length as preserved 4.4 m m (e x c l u d i n g a n t e n n a e a n d o v i p o s i t o r); fore wing 2.7 mm long; integument dark brown except metasoma lighter; body largely glabrous except legs bearing short, thick setae; integument without pronounced sculpturing or punctation, largely finely imbricate; propodeum coarsely and strongly areolate; wings clear and hyaline, veins brown to light brown in preserved colour. Head slightly shorter than wide in frontal view, width 0.95 mm and length 0.87 mm; genal width 0.2 mm; compound eyes ovoid, without prominent circumocular carina, 0.51 mm long, 0.3 mm wide; ocelli present and situated in a small triangle on top of vertex above compound eyes and immediately behind facial prominence, ocelli separated by approximately their diameter and from posterior border of head by about twice their diameter. Prominent facial horn projecting to a length of 0.37 mm in profile (from anterior toruli margin to apex of the horn). Antenna with scape ~3.5 times pedicel length, 0.43 mm long, 0.07 mm wide; pedicel more than twice as long as wide; flagellomeres each longer than wide except two apical most; flagellomere I the longest (0.31 mm long). Clypeal base indistinct from frons (epistomal sulcus lacking), with strong medial longitudinal ridge, ridge extends posteriorly with a small sharp tooth and anteriorly projects as small triangular prominence between mandibles. Palp formula (maxillary–labial) at least 6-3. Mandible massive, square shaped in frontal view, with straight outer and apical margins meeting in a rounded orthogonal angle; inner margin with ridges demarcating blunt ‘teeth’, apical tooth well projected without overlapping opposing mandible. Mesosoma laterally compressed, longer than high, length 2.2 mm, height 1.41 mm; pronotum prominent, with well-developed concave anterior surface, with poorly defined raised posterior surface, medially longer than wide, lateral surfaces large and slightly depressed; propleura long but not projecting anterior to pronotum; mesoscutum 0.53 mm long, notauli distinct; mesoscutellum convex in profile, 0.27 mm in its medial length; mesopleuron with small, oblique row of faint areolae demarcating border with pronotal lateral surface and more distinct, single row of areolae along border with metepisternum; propodeum weakly areolate, with prominent dorsal surface, dorsal length between metanotum and articulation with petiole equal to 0.3 mm. Legs long, with slender tarsi; meso- and metacoxae closer to each other than to procoxa; metafemur 1.81 mm long, metatibia 1.71 mm long; metabasitarsus elongate but slightly shorter than combined lengths of remaining tarsomeres, metabasitarsus length 0.8 mm, lengths of remaining tarsomeres (from base to apex), 0.34, 0.3, 0.16 and 0.23 mm. Formula of tibial spurs 1-2-2. Apex of all tarsi with a plantula. Outer pretarsal claw of foreleg tridentate (bearing two small pre-apical teeth), all other claws bidentate (with a single pre-apical tooth). Fore wing with costal space apically slightly broader than pterostigma; pterostigma longer than wide, tapering gradually in width to acute apex, margin inside marginal cell faintly convex; vein M+Cu forking after Sc+R midlength; 1M straight, nearly aligned with and longer than 1Rs; 1Rs originating prior to pterostigma a distance inferior to its length; 1rs-m meeting Rs slightly before r-rs; 2Rs longer than 1Rs; 2M small, meeting 1m-cu, then M abscissa extending toward wing apex; 2Rs slightly curved; r-rs originating from basal third of pterostigmal length, elongate, softly sinuous, as long as 2Rs; marginal cell broad, three time as long as wide, greatest width near tangent of pterostigmal apex; 3Rs mostly straight for majority of marginal cell length; two rs-m cross-veins present; second submarginal cell about as long as third and slightly widening towards the wing margin; third submarginal cell slightly wider posteriorly than anteriorly; first discal cell almost forming strongly slanted rhomboid (not distinctly pentagonal owing to exceptionally short 2M); 1m-cu straight and subparallel to 1M; cu-a originating at M+Cu fork, nearly aligned with 1M; 2m-cu present but weaker than surrounding veins; A2 present and meeting Cu apically in a small 2cu-a vein (demarcating second subbasal cell); vein 2A present. Hind wing with venation nearly complete, lacking only C; R with six distal hamuli and not meeting 2Rs apically; rs-m oblique, about twice as long as 1Rs, shorter than first free abscissa M; Cu+cu-a present, nearly orthogonal to M+Cu; apical abscissa Cu not visible if present; jugal lobe lacking. Metasoma with first segment forming tubular petiole 0.57 mm long, tergum and sternum fused without apparent indication of individual sclerites; gaster 1.72 mm long, compressed laterally, ovoid in profile, obviously longer than high, maximum height 0.84 mm, widening to the third gastral segment and then narrowing toward apex; ovipositor exerted, ≥ 1 mm long (not totally visible), with five minute teeth present on apical declivitous face; sheaths slightly longer than ovipositor. Male unknown.

KERATODELLITHA KIRIN MARÉCHAL, JOUAULT & PERRICHOT SP. NOV. (FIGS 5, 6) 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: 47865DDE-514F-4583-AB69-FF0D02A3BD5D Holotype: IGR.BU-021, preserved in rectangular piece of amber measuring 9 mm × 6 mm × 3 mm. Type locality and horizon: Hkamti site, Hkamti District, Sagaing Region, Myanmar; early Albian (~110 Mya), Early Cretaceous. Etymology: Named in reference to the fabled animal from Chinese mythology, Qilin or Kirin (DZDz). It was a chimeric creature that combined the traits of several animals and is a general allusion to the fierce and enigmatic habitus of the species. The specific epithet is a noun in apposition. Diagnosis: Scape short, about twice as long as pedicel; pretarsal claws of all legs bidentate; fore wing vein Rs+M longer than 2Rs; second and third submarginal cells equal in length; 1rs-m meeting Rs slightly after r-rs; 2rs-m complete and tubular; second discal cell short (about twice as long as wide); hind wing with five hamuli; metasoma with cylindrical gaster. Description: Female. Total length as preserved 4.87 mm (excluding antennae and ovipositor); fore wing ≥ 2.35 mm long; hind wing ~ 1.5 mm long; integument largely dark brown or clear brown, glabrous mesosoma without pronounced sculpturing or punctation except on propodeum, which is coarsely and strongly areolate; wings clear and hyaline, veins brown to light brown. Head wider than long, 0.64 mm wide; genal space ~ 0.2 mm; compound eye 0.42 mm long, 0.30 mm wide; three ocelli arranged in an equilateral triangle on vertex, ~ 0.05 mm in diameter each and distant from each other by 2.5 times their diameter; prominent facial horn projecting 0.44 mm in length (measured at apex); scape longer than wide, ~ 0.24 mm long; pedicel ~ 0.13 mm long, conical and shorter than flagellomere I; flagellomere I the longest, ~ 0.30 mm long; clypeal area partly hidden by debris within amber matrix; mandibles massive, rectangular, ~ 0.15 mm wide, the masticatory margin with large, blunt apical tooth followed by two(?) blunt teeth; outer margin of mandible covered by setae; palp formula (maxillary– labial) at least 5-3. Mesosoma laterally compressed, thinner than head, longer than high, 1.4 mm long, 0.94 mm high; pronotum well developed, 0.28 mm in maximal length; mesoscutum 0.55 mm long, slightly arched anteriorly; notauli not visible owing to preservation; mesopleuron wide, ~ 0.80 mm in maximal length, with small, oblique row of weak areolae along pronotal margin, and even weaker row of areolae along metepisternal margin; propodeum coarsely and strongly areolate, distinctly setose around petiolar insertion; length of propodeal dorsal surface between metanotum and articulation with metasomal petiole ~ 0.30 mm; legs long, thin, with five thin tarsomeres; meso- and metacoxae closer to each other than to procoxa; metacoxa 0.67 mm long, metatrochanter 0.29 mm long, metafemur 1.49 mm long, metatibia 1.29 mm long; metabasitarsus elongate, combined length of tarsomeres ~ 1.65 mm; tibial spurs formula 1-2-2, protibial spur bifid; pretarsal claws ~ 0.05 mm long, each with a single pre-apical tooth situated at about two-thirds of claw length. Fore wing with costal space apically about as broad as pterostigma; pterostigma much longer than wide, tapering gradually in width; marginal cell narrow, with vein 1Rs slightly convex; vein M+Cu forking distally at midlength of Sc+R; 1M straight to slightly curved, forming faint angle at junction with 1Rs; 1M shorter than 1Rs; 1Rs originating basal to pterostigma at a distance subequal to pterostigmal width; 1rs-m meeting Rs slightly after r-rs; Rs+M nearly straight; 2Rs greatly longer than 2M but conspicuously shorter than Rs+M; 2M exceedingly short and directed posteriorly to meet lm-cu; r-rs originating slightly before pterostigmal midlength, elongate, longer than 2Rs; marginal cell long and wide; two rs-m crossveins present, both tubular; second submarginal cell with posterior border 0.66 times shorter than that of third submarginal cell; second and third submarginal cells subequal in length; discal cell almost forming strongly slanted rhomboid (not distinctly pentagonal owing to exceptionally short 2M); lm-cu slightly concave, slightly shorter than lCu; 2m-cu conspicuous and tubular; 2cu-a present, enclosing subbasal cell. Hind wing with venation nearly complete, lacking only C; R with five distal hamuli, not meeting 2Rs apically; rs-m oblique, longer than 1Rs; Cu orthogonal to M+Cu; jugal lobe absent. Metasoma ~ 2.50 mm long; first metasomal segment forming a tubular petiole (0.66 mm long) slightly widening posteriorly, with tergum I and sternum I fused without apparent indication of individual sclerites; gaster slightly ovoid, elongate and laterally compressed (nearly flat, maybe owing to conservation), broadest at about its midlength; six tergites visible, respectively 0.32, 0.40, 0.40, 0.48, 0.72 and 0.16 mm long; ventral base of apical tergite covering ovipositor insertion; ovipositor exerted, moderately long, ~ 0.90 mm long (not totally visible), sheaths slightly longer than ovipositor. Male unknown., Published as part of Jouault, Corentin, Maréchal, Arthur, Condamine, Fabien L., Wang, Bo, Nel, André, Legendre, Frédéric & Perrichot, Vincent, 2022, Including fossils in phylogeny: a glimpse into the evolution of the superfamily Evanioidea (Hymenoptera: Apocrita) under tip-dating and the fossilized birth-death process, pp. 1396-1423 in Zoological Journal of the Linnean Society 194 on pages 1408-1410, DOI: 10.1093/zoolinnean/zlab034, http://zenodo.org/record/6458537

KERATODELLITHA ANUBIS JOUAULT, MARÉCHAL, WANG & PERRICHOT 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: D217B456-4EEF-4EF7-87B9-A497EE9BCA66 Holotype: NIGP174739, preserved in a rectangular piece of amber measuring 5 mm × 5 mm × 4 mm. Type locality and horizon: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; upper Albian to lower Cenomanian, mid-Cretaceous. Etymology: Named in reference to the Egyptian jackal god of the afterlife, Anubis, protector of graves and the deceased. The specific epithet is a noun in apposition. Diagnosis: Scape short, about twice as long as pedicel; pretarsal claws of all legs bidentate; Rs+M slightly longer than 2Rs; third submarginal cell longer than second one; 1rs-m aligned with r-rs; 2rs-m not fully tubular; second discal cell long, about three times as long as wide; hind wing with five hamuli; metasoma with cylindrical gaster. Description: Male. Total length as preserved 4.06 mm (excluding antennae and ovipositor); fore wing length ≥ 2.2 mm; integument largely dark brown and largely glabrous; integument without pronounced sculpturing or punctation, only finely imbricate; propodeum coarsely and strongly areolate; wings clear and hyaline, veins brown to light brown in preserved colour. Head wider than long; clypeus and frons vertical and flat; genal width at about midlength 0.1 mm; compound eye 0.41 mm long, 0.27 mm wide; ocelli protruding, large, arranged in small triangle on top of vertex above compound eyes, separated by approximately their diameter and from posterior border of head by twice their diameter. Prominent facial horn projecting to length of 0.25 mm in profile (from anterior toruli margin to apex of horn); scape longer than wide, twice as long as pedicel, 0.2 mm long, 0.07 mm wide; pedicel more than twice as long as wide; flagellomere I the longest (~ 0.25 mm long). Clypeal base indistinct from frons, without medial longitudinal ridge. Palpal formula at least 5-3. Mandible large, square in frontal view, with straight outer and apical margins, margins meeting at rounded obtuse outer angle; inner margin with four blunt teeth and with one stout and longer, pointed apical tooth. Mesosoma laterally compressed, longer than high, 1.35 mm long, ≥ 0.8 mm high; pronotum well developed, with well-developed concave anterior surface, apparently slightly longer than wide, lateral surfaces large and slightly depressed; mesoscutum with weak notauli not meeting, mesoscutum 0.5 mm long; mesoscutellum well separated from mesoscutum, slightly convex in profile, 0.2 mm long; mesopleuron with oblique depression and small, single row of weak areolae demarcating border with pronotal lateral surface, and even weaker row of areolae along border with metepisternum; propodeum coarsely and strongly areolate, dorsal length between metanotum and articulation with petiole equal to 0.23 mm. Legs not fully preserved except right hindlegs; long, with meso- and metacoxae closer than procoxa; hind femur medially swollen. Fore wing with costal space apically about as broad as pterostigma; pterostigma much longer than wide, tapering gradually in width in apical half to acute apex; marginal cell narrow, with vein 1Rs slightly convex; vein M+Cu forking distally to midlength of Sc+R; 1M slightly curved, forming faint angle at junction with 1Rs, 1M subequal to 1Rs; 1Rs originating basal to pterostigma at distance subequal to pterostigmal width; 1rs-m aligned with r-rs; Rs+M straight; 2Rs much longer than 2M but slightly shorter than Rs+M; 2M exceedingly short and directed posteriorly to meet lm-cu; 3M slightly convex basally, then longitudinal; 2Rs slightly curved near Rs+M, then almost straight; r-rs originating before pterostigmal midlength, weakly sinuous, elongate, slightly shorter than 2Rs; marginal cell damaged and partial; two rs-m cross-veins present, second one distinctly weaker and thinner than surrounding veins (but not nebulous); second submarginal cell with posterior border 0.7 times shorter than posterior border of third one; third submarginal cell ~1.3 times longer than second; discal cell almost forming strongly slanted rhomboid (not distinctly pentagonal owing to exceptionally short 2M), lm-cu straight, 1.3 times longer than lCu; lCu-a slightly distal with 1M; 2m-cu conspicuous and tubular; 2cu-a present, enclosing subdiscal cell. Hind wing with venation nearly complete except lacking C; R with five distal hamuli, not meeting 2Rs apically; rs-m oblique, longer than 1Rs, about twice as short as first 1M; Cu present, virtually orthogonal to M + Cu; jugal lobe lacking. Metasoma 2.0 mm long; first segment forming trapezoidal petiole, narrower basally, 0.44 mm long; tergum I and sternum I fused, without apparent indication of individual sclerites; gaster compressed laterally, gradually and slightly tapering in height from midlength to apical segment. Parameres directed backward, keel shaped, protruding ~ 0.2 mm. Aedegus not clearly visible. Female unknown., Published as part of Jouault, Corentin, Maréchal, Arthur, Condamine, Fabien L., Wang, Bo, Nel, André, Legendre, Frédéric & Perrichot, Vincent, 2022, Including fossils in phylogeny: a glimpse into the evolution of the superfamily Evanioidea (Hymenoptera: Apocrita) under tip-dating and the fossilized birth-death process, pp. 1396-1423 in Zoological Journal of the Linnean Society 194 on pages 1405-1408, DOI: 10.1093/zoolinnean/zlab034, http://zenodo.org/record/6458537

KEY TO SPECIES OF KERATODELLITHA The three species of Keratodellitha can be distinguished mainly by the characters given in the key below. 1. Scape short (about twice as long as pedicel); fore wing with Rs+M longer than 2Rs; hind wing with five hamuli; metasoma cylindrical............................................................................................................................ 2 Scape long (more than three times as long as pedicel); fore wing with Rs+M as long as 2Rs; hind wing with six hamuli; metasoma rounded................................................................................. Keratodellitha basilisci 2. Fore wing with third and second submarginal cells similar in length, second discal cell about twice as long as wide............................................................................................................................. Keratodellitha kirin Fore wing with third submarginal cell longer than second one, second discal cell conspicuously longer than wide (three times)......................................................................................................... Keratodellitha anubis, Published as part of Jouault, Corentin, Maréchal, Arthur, Condamine, Fabien L., Wang, Bo, Nel, André, Legendre, Frédéric & Perrichot, Vincent, 2022, Including fossils in phylogeny: a glimpse into the evolution of the superfamily Evanioidea (Hymenoptera: Apocrita) under tip-dating and the fossilized birth-death process, pp. 1396-1423 in Zoological Journal of the Linnean Society 194 on page 1412, DOI: 10.1093/zoolinnean/zlab034, http://zenodo.org/record/6458537