Corentin Jouault
email: jouaultc0@gmail.com

Arthur Maréchal
email: marechal.arthur.pro@gmail.com

Fabien L Condamine
email: fabien.condamine@gmail.com

Bo Wang
email: bowang@nigpas.ac.cn

André Nel
email: anel@mnhn.fr

Frédéric Legendre
email: frederic.legendre@mnhn.fr

Vincent Perrichot
email: vincent.perrichot@univ-rennes1.fr

Including fossils in phylogeny: A glimpse into the evolution of the superfamily Evanioidea (Hymenoptera, Apocrita) under tip-dating and the fossilized birth-death process

Keywords: Keradellitha anubis Jouault, Bayesian inferences, fossilized birth-death, morphological phylogeny, time divergence, calibration

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INTRODUCTION

Recent advances in phylogenomics have clarified relationships within the hymenopteran lineages and shown the monophyly of all superfamilies except Vespoidea [START_REF] Sharkey | Phylogenetic relationships among superfamilies of Hymenoptera[END_REF]Ronquist et al., 2012a;[START_REF] Peters | Evolutionary history of the Hymenoptera[END_REF][START_REF] Sharanowski | Phylogenomics of Ichneumonoidea (Hymenoptera) and implications for evolution of mode of parasitism and viral endogenization[END_REF]. Nonetheless, the monophylies of these clades are not so clear-cut when fossils are considered. The most recent works have estimated robust time-calibrated phylogenies for Hymenoptera based on analyses of genomic data, and have found evidence that Evanioidea is the sister lineage of Stephanoidea [START_REF] Peters | Evolutionary history of the Hymenoptera[END_REF][START_REF] Tang | Mitochondrial phylogenomics of the Hymenoptera[END_REF]. Additionally, molecular and morphological studies strongly support the monophyly of Evanioidea [START_REF] Dowton | Molecular phylogeny of the insect order Hymenoptera: apocritan relationships[END_REF][START_REF] Dowton | Molecular phylogeny of the apocritan wasps: the Proctotrupomorpha and Evaniomorpha[END_REF][START_REF] Vilhelmsen | Beyond the wasp waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera)[END_REF][START_REF] Heraty | Evolution of the hymenopteran megaradiation[END_REF][START_REF] Sharkey | Phylogenetic relationships among superfamilies of Hymenoptera[END_REF][START_REF] Klopfstein | The hymenopteran tree of life: evidence from protein-coding genes and objectively aligned ribosomal data[END_REF][START_REF] Payne | Direct optimization, sensitivity analysis, and the evolution of the hymenopteran superfamilies[END_REF][START_REF] Li | Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar[END_REF][START_REF] Sharanowski | Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera)[END_REF]. The superfamily Evanioidea is currently composed of five fossil families ( †Praeaulacidae, †Anomopterellidae, †Andreneliidae, †Baissidae, and †Othniodellithidae) that are widely represented in Mesozoic deposits, plus three extant families (Evaniidae, Aulacidae, Gasteruptiidae) with a rather extensive fossil record. The Evanioidea may have arisen during the Late Triassic [START_REF] Sharkey | Phylogenetic relationships among superfamilies of Hymenoptera[END_REF][START_REF] Peters | Evolutionary history of the Hymenoptera[END_REF][START_REF] Tang | Mitochondrial phylogenomics of the Hymenoptera[END_REF] but the evolutionary history and the molecular age estimates within the superfamily have not yet been studied, except for its extant families [START_REF] Sharanowski | Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera)[END_REF]Parslow et al., 2020a).

Extant evanioid wasps are known from 310 aulacid, 580 evaniid, and about 500 gasteruptiid species (see details in [START_REF] Ramage | New species and records of Evanioidea and Stephanoidea from New Caledonia (Hymenoptera)[END_REF]) but data on their biology are limited. Several studies report aulacid wasps as koinobiont endoparasitoids of wood-boring larvae of Hymenoptera Xiphydriidae and Coleoptera Cerambycidae and Buprestidae (e.g. [START_REF] Deyrup | A maple wood wasp, Xiphydria maculata, and its insect enemies (Hymenoptera: Xiphydriidae)[END_REF][START_REF] Gauld | The evaniomorph parasitoid families[END_REF][START_REF] Jennings | Biology and host relationships of aulacid and gasteruptiid wasps (Hymenoptera: Evanioidea): a review[END_REF][START_REF] Kuroda | A revisional study of the genus Aulacus Jurine (Hymenoptera: Aulacidae) of Japan[END_REF]. Adults of gasteruptiids apparently feed on nectar and pollen [START_REF] Jennings | Biology and host relationships of aulacid and gasteruptiid wasps (Hymenoptera: Evanioidea): a review[END_REF], while their larvae are predator-inquiline, feeding on larvae or larval food of solitary bees (Antophila of the families Apidae, Colletidae, Halictidae, known from numerous specimens in the highly fossiliferous Burmese amber, representing respectively 2.4% and 5.2% of the valid species (Jouault et al., 2021b). Thus it seems that the dominance of the ichneumonoids occurred during the middle to Late Cretaceous even if the evanioids are already abundant during this period (Tabl. 1).

In this study we present and describe three new species of othniodellithid wasps belonging to a new genus. We also revise the fossil record of Evanioidea and propose a time-calibrated phylogeny for the group using a tip-dating approach. The analysis of the fossil record of the superfamilies Evanioidea and Ichneumonoidea is used to document the shift in specific richness between Evanioidea and Ichneumonoidea in the Cretaceous. We also discuss the record of two species belonging to a same genus but originating from two deposits of different ages.

MATERIAL AND METHODS

SPECIMEN SAMPLING AND MORPHOLOGICAL EXAMINATION

The two dark orange amber pieces containing Keradellitha basilici Jouault, Maréchal, Wang & Perrichot gen. et sp. nov. and K. anubis Jouault, Maréchal, Wang & Perrichot sp. nov. originate from the deposits of Noije Bum in the Hukawng Valley (26° 29' N, 96° 35' E), Kachin State, Northern Myanmar (see Grimaldi & Ross, 2017: fig 2). Radiometric data established an early Cenomanian age (98.79 ± 0.62 Ma) for Kachin amber, based on zircons from volcanic clasts found within the amber-bearing sediments [START_REF] Shi | Age constraint on Burmese amber based on U-Pb dating of zircons[END_REF]. Some ammonites found in the amber-bearing bed and within amber corroborates a late Albian / early Cenomanian age [START_REF] Cruickshank | Geology of an amber locality in the Hukawng Valley, northern Myanmar[END_REF][START_REF] Yu | An ammonite trapped in Burmese amber[END_REF]. The amber piece, also dark orange, containing Keradellitha kirina Maréchal, Jouault & Perrichot sp. nov., originates from the Hkamti site (about 80 km southwest of the Angbamo site), Hkamti District, Sagaing Region, Myanmar; see detailed map in Zheng et al. (2018: supplementary figure 2) or in Xing & Qiu, (2020: fig 1). An early Albian age (109.7 ± 0.4 Ma) was established for Hkamti amber based on zircon U-Pb analyses of clastic sediments [START_REF] Xing | Zircon U-Pb age constraints on the mid-Cretaceous Hkamti amber biota in northern Myanmar[END_REF].

Maximum parsimony (MP) analyses of the morphological dataset (Appendix S1) were conducted with PAUP 4.0a166 [START_REF] Swofford | PAUP: Phylogenetic Analysis Using Parsimony (and Other Methods)[END_REF]. Outgroup taxa were treated as paraphyletic with respect to the ingroup. Tree searches were performed using a heuristic search method with the following options: maximum number of trees saved equal to 10,000, only optimal trees retained, collapse of zero-length branches, and a tree bisection and reconnection (TBR) swapping algorithm. When searches produced more than one optimal cladogram, a strict consensus was performed (Suppl. Fig. 1). To measure the robustness of the parsimony cladograms, bootstrap analyses [START_REF] Felsenstein | Confidence limits on phylogenies: an approach using the bootstrap[END_REF][START_REF] Hillis | An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis[END_REF] were executed using the full heuristic search option for 100 replicates. We considered values of bootstrap support (BS) equal or above 70 as strong node supports [START_REF] Hillis | An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis[END_REF].

BAYESIAN PHYLOGENETIC INFERENCE AND DIVERGENCE TIME ESTIMATES

We carried out Bayesian inferences (BI) using a relaxed clock model in MrBayes 3.2.7a [START_REF] Huelsenbeck | MRBAYES: Bayesian inference of phylogeny[END_REF][START_REF] Ronquist | MrBayes 3: Bayesian phylogenetic inference under mixed models[END_REF]Ronquist et al., 2012b). We performed tip-dating analyses with a fossilized birth-death (FBD) model wherein fossil taxa are terminals. Analyses were computed with a Markov one parameter (Mkv) model [START_REF] Lewis | A likelihood approach to estimating phylogeny from discrete morphological character data[END_REF] with or without a gamma rate variation across characters and with the independent gamma relaxed clock model [START_REF] Lepage | A general comparison of relaxed molecular clock models[END_REF]lset rates = invgamma, prset clockvarpr = igr, prset igrvarpr = exp(10); no rate variation was computed with lset rates = equal). Following Ronquist et al. (2012a), the prior used for the mean clock rate was gamma (2, 200), except in one analysis wherein we tested a normal but very flat prior (prset clockratepr = normal(0.0025,1) and a flat uniform speciation prior (prset speciationpr = uniform(0,10); [START_REF] Matzke | Inferring node dates from tip dates in fossil Canidae: the importance of tree priors[END_REF]. The proportion of extant taxa was set to 0.015 (21 extant species out of ca. 1,400 species). Sampling strategy of taxa was either set to diversity (prset samplestrat = diversity wherein fossils are sampled randomly and can be tips or ancestors), random (prset samplestrat = random wherein fossil can be tips or ancestors) or fossil tip (prset samplestrat = fossiltip wherein fossil and taxa are assumed to be sampled randomly and fossil ancestor is not allowed). An exponential prior and a beta prior were used for the net speciation rate and the relative extinction rate using the following functions: prset speciationpr = exp(100), and prset extinctionpr = beta (1,1), respectively. In all our tip-dating analyses, the node age prior was set to 'calibrated'. All analyses comprised two runs and four Markov chains Monte Carlo (MCMC), and were launched for 50 million generations. MCMC were sampled every 5000 generations and a burn-in fraction of 0.25 was used. Convergence diagnostics were checked for each analysis with the average standard deviation of split frequencies < 0.01, PRSF close to 1.0 in MrBayes outputs, and ESS > 200 in Tracer 1.7.1 (Rambaut et al., 2018). We performed two tip-dating analyses: (1) we used extinct taxa calibrated with uniform distributions bounded according to the minimum and maximum ages of their deposits [START_REF] Barido-Sottani | Ignoring stratigraphic age uncertainty leads to erroneous estimates of species divergence times under the fossilized birth-death process[END_REF]Appendix S2), and (2) we used extinct taxa calibrated with fixed distributions bounded according to the minimum ages of their deposits. In the FBD model, all fossils were set as tips (r = 1). The prior probability distribution on branch lengths was set to clock:fossilization. Three nodes were assigned offset exponential priors: (1) the root of the tree (minimum age: 183 Ma, mean age: 245 Ma), (2) the clade Apocrita (183, 236 Ma), and (3) the ingroup (164 Ma, 178 Ma). Those ages were set according to the latest studies (Ronquist et al., 2012a;[START_REF] Peters | Evolutionary history of the Hymenoptera[END_REF][START_REF] Tang | Mitochondrial phylogenomics of the Hymenoptera[END_REF]. All scripts are provided as Supporting Information. Etymology. The new generic name is a Latinised combination of the Greek words, kéras ('horn') and dellithos ('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 circum-ocular carina, inner lateral margins slightly concave; toruli widely separated by cephalic horn, located on lateral surfaces, opening dorso-laterally; 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).

KERADELLITHA BASILICI Jouault, Maréchal, Wang & Perrichot, sp. nov. urn:lsid:zoobank.org:act:C67E5240-F1BC-4836-86F7-3B9D1D375AA8 (Figs. [1][2] Holotype: NIGP174738, preserved in a rectangular piece of amber measuring 17 × 10 × 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 a mythical beast (Basilicus) 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 to be treated as a noun in a genitive case.

Diagnosis. Scape long, about 3,5 times longer than pedicel; outer (external) pretarsal claw of fore leg with two pre-apical 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), roundedshape.

Description. Female. Total length as preserved 4.4 mm (excluding antennae and ovipositor); 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 color.

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 circum-ocular 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 length of 0.37 mm in profile (from anterior toruli margin to apex of the horn). Antenna with scape about 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, squareshaped 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 long, 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 meta-coxae 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 mm, 0.3 mm, 0.16 mm, and 0.23 mm. Tibial spurs formula 1-2-2. Apex of all tarsi with a plantula.

Outer pretarsal claw of fore leg 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 mid-length; 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 rrs; 2Rs longer than 1Rs; 2M very small, meeting 1m-cu, then M abscissa extend toward wing apex; 2Rs slightly curved; r-rs originating from basal third of pterostigmal length, elongate, softly sinu-ous, 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 crossveins 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 sub-parallel to 1M; cu-a originating at M+Cu fork, nearly aligned with 1M; 2mcu present but weaker than surrounding veins; A2 present and meeting Cu apically in a small 2cu-a vein (demarcating second sub-basal cell); vein 2A present. Hind wing with venation nearly complete, only lacking 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, at least 1 mm long (not totally visible), with five minute teeth present on apical declivitous face; sheaths slightly longer than ovipositor. Etymology. Named in reference to the mythological Egyptian god Anubis, protector of graves and the deceased. The specific epithet 'anubis' is to be treated as a noun in apposition.

Male unknown.

KERADELLITHA ANUBIS

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 at least 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 color.

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 (about 0.25mm 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, at least 0.8mm 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 hind legs; long, with meso-and meta-coxae closer than pro-coxa; 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 crossveins 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 about 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. Etymology. Named in reference to the fabulous animal from Chinese mythology: Qilin or Kirin 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 to be treated as a noun in a genitive case.

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 at least 2.35 mm long; hind wing ca. 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 about 0.2 mm; compound eye 0.42 mm long, 0.30 mm wide; three ocelli arranged in an equilateral triangle on vertex, about 0.05 mm in diameter each and distant from each other by 2.5 times their diameter; prominent facial horn projecting for 0.44 mm long (measured at apex); scape longer than wide, about 0.24 mm long; pedicel about 0,13 mm long, conical and shorter than flagellomere I; flagellomere I the longest, about 0.30 mm long; clypeal area partially hidden by debris within amber matrix; mandibles massive, rectangular, about 0.15 mm wide, the masticatory margin with large, blunt apical tooth followed by 2(?) 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 due to preservation; mesopleuron wide, ca. 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 about 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 ca. 1.65 mm long; tibial spurs formula 1-2-2, protibial spur bifid; pretarsal claws ca. 0.05 mm long, each with a single pre-apical tooth situated at about 2/3 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 of 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 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 sub-basal cell. Hind wing with venation nearly complete, only lacking 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 about 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 due to conservation), broadest at about its midlength; six tergites visible, respectively 0.32 mm, 0.40 mm, 0.40 mm, 0.48 mm, 0.72 mm, and 0.16 mm long; ventral base of apical tergite covering ovipositor insertion; ovipositor exerted, moderately long, ca. 0.90 mm long (not totally visible), sheaths slightly longer than ovipositor.

Male unknown.

RESULTS

We summarized the fossil diversity of Evanioidea and Ichneumonoidea (Fig. 7) using the function count taxa in Paleobiology Database (http://fossilworks.org; last access November 25, 2020). Although diversity changes through time as estimated by this analysis are biased by the incompleteness of the fossil record, we could extract one general pattern showing that Evanioidea were already diversified in the Jurassic period with more than 70 species (Fig. 7), while the earliest record of Ichneumonoidea (Cretobraconus maculatus [START_REF] Rasnitsyn | New Eoichneumonidae from Early Cretaceous of Siberia and Mongolia (Hymenoptera: Ichneumonoidea)[END_REF] was described from the Lower Cretaceous Ulugei Formation [START_REF] Rasnitsyn | New Eoichneumonidae from Early Cretaceous of Siberia and Mongolia (Hymenoptera: Ichneumonoidea)[END_REF][START_REF] Kopylov | The Khasurty fossil insect lagerstatte[END_REF].

Afterwards, the species number of Ichneumonoidea increased, until becoming almost equivalent to that of Evanioidea during the Early Cretaceous. After the Cretaceous, the number of species of Ichneumonoidea clearly exceeded the number of Evanioidea species with a peak during the Eocene (Fig. 7).

Adding 41 species to the morphological matrix has provided a relatively strong support for the monophyly of Evanioidea (PP = 1; BS = 63). The resulting phylogeny has also clarified the rela -tionships among and between extant and extinct taxa (Fig. 8; Supplementary Fig. 1). We found the genus Hyptiogastrites as an early-diverging lineage within the Aulacidae, and Vectevania vetula [START_REF] Cockerell | Two families of insects new to British Tertiary strata[END_REF] among the Aulacidae as an early-diverging taxon relative to the crown-Aulacidae (Aulacus + Pristaulacus). Although weakly supported, the Nevaniinae were recovered as a monophyletic subfamily within the Praeaulacidae (PP = 0.61; BS = 68), and sister lineage of the genus Eosaulacus (PP = 0.13). The two species of Keradellitha gen. nov. coded in the matrix were found nested within the †Othniodellithidae in BI and MP, confirming our taxonomic attributions (Fig. 8; Supplementary Fig. 1). We summarized the phyletic states of each family in Tabl. 2. In all our Bayesian analyses, Evanioidea, †Anomopterellidae (PP = 0.93; BS = 72), Gasteruptiidae (PP = 0.91; BS = 66), †Othniodellithidae (PP = 1; BS = 99), †Andreneliidae, Aulacidae (PP = 0.67), and Evaniidae (PP = 0.99; BS = 50) were recovered as monophyletic lineages, while †Praeaulacidae (PP = 0.24) and †Baissidae were recovered as paraphyletic. In the MP analysis, the family Aulacidae was polyphyletic (Tabl. 2; Supplementary Fig. 1). Based on these results, we proposed a new hierarchical suprageneric classification of Evanioidea (Tabl. 3).

Following our modified suprageneric classification of Evanioidea, the 'protoevanioides' clade ( †Praeaulacidae + †Othniodellithidae) (PP = 0.24) corresponds to the earliest radiation of the evanioid wasps, characterized by their similarities shared with †Ephialtitidae. The second and main radiation of evanioid wasps, the 'neoevanioides' clade (PP = 0.95), comprises the families †Anomopterellidae, †Baissidae, Gasteruptiidae, †Andreneliidae, Aulacidae, and Evaniidae.

Besides the relationships of the new fossil taxa, we estimated divergence dates within Evanioidea (Fig. 8). Using tip-dating analyses (run with the Mkv+G, FBD model, uniform distribution, and samplestrat = fossiltip), we estimated the divergences of stem-and crown-Evanioidea to the Late Triassic and Early Jurassic: 203 Ma (95% HPD = 188-220 Ma) and 176 Ma (95% HPD = 159-194 Ma), respectively. The tip-dating analysis also indicated that the crownprotoevanioides originated around 201 Ma (95% HPD = 186-215 Ma). Among 'protoevanioides', †Othniodellithidae were estimated to have diverged from their Praeaulacidae ancestor about 117 Ma (95% HPD = 110-127 Ma). The crown-'neoevanioides' was estimated at 190 Ma (95% HPD = 175-208 Ma). Among neoevanioides, we have also estimated the crown-euaulacides (Aulacidae + Gasteruptiidae) around 139 Ma (95% HPD = 121-158 Ma). Aulacidae were recovered as sister lineage to Gasteruptiidae and their crown-groups would date back to 68 Ma (95% HPD = 52-89 Ma) and 20 Ma (95% HPD = 6-36 Ma), respectively. Extant Evaniidae have a crown age at 58 Ma (95% HPD = 42-75 Ma), while their oldest relatives would date back to the Middle Jurassic ( 155Ma, 95% HPD = 142-168 Ma). Divergence time estimates for the main evanioid lineages are summarized in Tabl. 4.

DISCUSSION

SYSTEMATIC PLACEMENT OF NEW TAXA

Our fossils present all the diagnostic characters of the family †Othniodellithidae as described by Engel et al. (2016a) and emended by [START_REF] Engel | New evanioid wasps from the Cenomanian of Myanmar (Hymenoptera: Othniodellithidae, Aulacidae), with a summary of family-group names among Evanioidea[END_REF], including the complete wing venation with two rs-m crossveins and 2m-cu usually present (see Engel et al., 2016a: fig 3); vein M+Cu forking distad midlength of Sc+R; vein lcu-a confluent to slightly distal to 1M; marginal cell large, wide near tangent with apex of pterostigma and elongate, nearly reaching to wing apex. With a nearly complete hind wing venation, the fossils may be confused with †Praeaulacidae, except that othniodellithids are lacking the vein C and a jugal lobe. Also, our new taxa cannot be assigned to †Praeaulacidae as they have fewer antennomeres (22 vs. at least 31 in Praeaulacidae). Overall, they possess the typical horn of the †Othniodellithidae (synapomorphy); the same wing venation (with minor variations in the presence of vein 2m-cu or alignment of vein M+Cu and Cu); and the typical habitus of the family with well-developed pronotum and mesoscutum, and a metasoma separated from mesosoma with a petiole. by the antenna with a short scape (vs. long); the fore wing Rs+M longer than 2Rs (vs. equal), third submarginal cell longer than the second one (vs. equal), subdiscal cell long (vs. short and wide), 2rs-m incomplete (vs. complete and tubular); the hind wing with five hamuli (vs. six); and its metasoma cylindric (vs. rounded). For additional comparisons, see the keys proposed above.

COMPARISONS OF DIVERGENCE TIME ESTIMATES FOR EVANIOIDEA

As previously mentioned, divergence time estimates have only been proposed for the superfamily Evanioidea, including the Aulacidae, Evaniidae, and Gasteruptidae. Our result from the tipdating approach of the superfamily Evanioidea is congruent with previous studies (Ronquist et al., 2012a;[START_REF] Peters | Evolutionary history of the Hymenoptera[END_REF][START_REF] Li | Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar[END_REF][START_REF] Sharanowski | Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera)[END_REF][START_REF] Tang | Mitochondrial phylogenomics of the Hymenoptera[END_REF], which estimated the origin of Evanioidea in the Middle Jurassic, or possibly the Late Triassic. A recent paper has proposed the divergence of Ma, with lognormal prior) and around 151.5 Ma (135.9-166.7 Ma, with normal prior) [START_REF] Sharanowski | Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera)[END_REF]. Our estimates are congruent with these results since our minimum and maximum time estimates range between 140 and 186 Ma. Based on our analyses, the first known diversification of the stem-Aulacidae is estimated during the Early Cretaceous (around 130 Ma, Fig. 8), which would be earlier than the first known diversification of the stem-Gasteruptiidae, estimated in the mid-Cretaceous (around 105 Ma, Tabl. 4, Fig. 8). Interestingly, the age of the first diversification of the Gasteruptiidae that we recovered corresponds to the oldest records of the Antophila that they predate [START_REF] Debevec | Identifying the sister group to the bees: a molecular phylogeny of Aculeata with an emphasis on the superfamily Apoidea[END_REF][START_REF] Cardinal | The evolution of floral sonication, a pollen foraging behavior used by bees (Anthophila)[END_REF][START_REF] Genise | 100 Ma sweat bee nests: early and rapid codiversification of crown bees and flowering plants[END_REF]. [START_REF] Sharanowski | Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera)[END_REF] proposed the divergence between Aulacidae and Gasteruptiidae around 111.6 Ma, which is likely underestimated because the earliest fossil record of aulacid wasp is dated from the mid-Cretaceous (Engel & Wang, 2016a). The most recent study investigating the Gasteruptiidae phylogeny estimated that the clade (Aulacidae + Gasteruptiidae) has diverged from Evaniidae ca. 151 Ma (Parslow et al., 2020a). This is younger than our divergence estimate ca. 176 Ma, congruent with the numerous Evaniidae known from the Early Cretaceous Lebanese amber (Tabl. 1). According to Parslow et al. (2020a), the divergence between Aulacidae and Gasteruptiidae occurred ca. 83 Ma, which is highly incongruent with the oldest representatives of both families both known from mid-Cretaceous Burmese amber. However, they estimated the crown-Aulacidae to arise ca. 52 Ma, close to the 68 Ma estimate in our analysis. They proposed that the crown-gasteruptiids arose ca. 60 Ma while we only recover them ca. 20 Ma. This latter difference is directly influenced by the scarcity of the gasteruptiid fossil record. As for the estimate of crown-Aulacidae, Parslow et al. (2020a) may have slightly underestimated the age of stem-Evaniidae (ca. 137 Ma vs. 163 Ma in our analyses). Alternatively, analyses with the FBD process might, in certain circumstances that still need to be clearly identified, overestimate datings [START_REF] Arcila | An evaluation of fossil tip-dating versus node-age calibrations in tetraodontiform fishes (Teleostei: Percomorphaceae)[END_REF][START_REF] O'reilly | Tips and nodes are complementary not competing approaches to the calibration of molecular clocks[END_REF][START_REF] Matschiner | Selective sampling of species and fossils influences age estimates under the Fossilized Birth-Death model[END_REF].

REFINING STEM AND CROWN-AGES WITHIN EVANIOIDEA

Recently, a hierarchical, suprageneric classification of Evanioidea was proposed without reconstruction of a phylogenetic hypothesis to support this classification (Engel, 2017: tabl. 2). However, the former classification was almost similar to the one derived from our Bayesian analyses (Engel, 2017: tabl. 2). Therefore, we only slightly modified it to fit with our phylogenetic results (Tabl. 2).

Contrary to many other hymenopteran taxa, the Evanioidea had a greater diversity (in terms of family) in the deep past than today. This peculiar diversity pattern implies the extinctions or declines of several lineages. The study of this fossil record allows differentiating stem-from crowngroups, which is key to our understanding of how lineages waxed and waned [START_REF] Budd | The dynamics of stem and crown groups[END_REF]. Since no phylogenetic analyses integrated both fossil and extant representatives in a timecalibrated phylogeny, the delimitation and age estimates of these groups were not clearly established in evanioids. Our study thus brings the first phylogeny-based attempt in this regard.

Among the 'Neoevanioides', the delineation of stem-and crown-groups has been challenged by the misplacement of controversial taxa, namely Hyptiogastrites [START_REF] Cockerell | Fossil insects[END_REF], Vectevania Cockerell, 1922, and †Andreneliidae. The monospecific genus Hyptiogastrites has been assigned to †Baissidae [START_REF] Li | Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar[END_REF]. However, following the description of a new male specimen [START_REF] Turrisi | New aulacid wasps from the mid-Cretaceous of Myanmar (Hymenoptera: Evanioidea)[END_REF] and its coding in the current phylogenetic matrix, it is now recovered in the Aulacidae. This placement is congruent with a detailed morphological study and cladistic analysis [START_REF] Jennings | Biology and host relationships of aulacid and gasteruptiid wasps (Hymenoptera: Evanioidea): a review[END_REF] but also with the former suprageneric classification proposing Hyptiogastrites in an early-diverging position within Aulacidae [START_REF] Engel | New evanioid wasps from the Cenomanian of Myanmar (Hymenoptera: Othniodellithidae, Aulacidae), with a summary of family-group names among Evanioidea[END_REF]. As a result, Hyptiogastrites should be transferred from †Baissidae to Aulacidae. Initially described from a single partially preserved specimen, from the latest Eocene Bembridge Marls in England [START_REF] Cockerell | Two families of insects new to British Tertiary strata[END_REF], Vectevania vetula [START_REF] Cockerell | Two families of insects new to British Tertiary strata[END_REF] was formerly ascribed to the Evaniidae. Recently, its position was revised [START_REF] Rasnitsyn | Vectevania vetula Cockerell, 1922 from the uppermost Eocene of Bembridge Marls, England, and the system of the family Gasteruptiidae s.l. [(Vespida (=Hymenoptera), Evanioidea[END_REF], and V. vetula was placed among the 'Aulacinae' (= Aulacidae). The most recent morphology-based phylogenetic analysis of the Evanioidea [START_REF] Li | Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar[END_REF] has led its authors to erect a monogeneric family for this taxon. It seems, however, unlikely that Vectevania belongs to a family distinct from the other evanioid families because Vectevaniidae was not supported by any putative autapomorphic trait (Li et al., 2018: fig 24). We recovered Vectevania vetula among the Aulacidae as an early-diverged taxon close to the extant genera (Aulacus + Pristaulacus).

The phylogenetic position of the family †Andreneliidae has long been controversial since Rasnitsyn (pers. comm. in Turrisi & Ellenberger, 2019) proposed the †Andreneliidae as an 'intermediate' clade between Evaniidae and (Aulacidae + Gasteruptiidae + †Baissidae) [START_REF] Turrisi | New aulacid wasps from the mid-Cretaceous of Myanmar (Hymenoptera: Evanioidea)[END_REF]. Here, the family †Andreneliidae occupies an early-diverging position relative to Evaniidae, consistent with the results of the recent morphology-based phylogeny [START_REF] Li | Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar[END_REF].

The challenging placement of the †Andreneliidae results from the poor preservation of the unique known andreneliid specimen, which does not allow confident discussion on its placement. In this way, we define the stem-and crown-groups of Gasteruptiidae, Aulacidae and Evaniidae as shown in Figure 8.

Therefore, we consider that the Evanioidea comprise eight families: †Andreneliidae, †Anomopterellidae, Aulacidae, †Baissidae, Evaniidae, Gasteruptiidae, †Othniodellithidae, and †Praeaulacidae.

OPPOSITE DIVERSITY PATTERN BETWEEN EVANIOIDEA AND ICHNEUMONOIDEA

A recent study provided new data about variations in the species richness of hymenopteran taxa, from the Early Cretaceous to present, and showed that the Evanioidea represented more than 20% of the hymenopteran species (at least 64 species) among non-aculeate apocritan wasps during the Early Cretaceous (Jouault et al., 2021b: fig 6). Based on these previous results, it appears that a shift in relative proportions between Evanioidea and Ichneumonoidea occurred during the Cretaceous. The evanioids flourished during the Jurassic while no ichneumonoids are known during this period. During the Early Cretaceous the proportion of evanioids and ichneumonoids varies and depends on the deposits: in the Khasurty deposits (Transbaikalian) the abundance of both superfamilies are similar [START_REF] Kopylov | The Khasurty fossil insect lagerstatte[END_REF], while the evanioids prevail in the Gurvan-Eren Formation (western Mongolia: Rasnitsyn, 1990a). On the contrary the ichneumonoids are more diverse in the Bon-Tsagan deposits (Mongolia: Rasnitsyn, 1980b). The relative species richness of ichneumonoids and evanioids among non-aculeates shifted from ca. 25% during the middle to Late Cretaceous to more than 50% (for ichneumonoids) vs. less than 5% (for evanioids) after the early Late Cretaceous (Jouault et al., 2021b: fig 6). This significant proportion of ichneumonoids almost invariably dominate the Late Cretaceous with an abundant ichneumonoid fauna described from the Yantardakh deposits (Taimyr) or from the Ola Formation (Rasnitsyn, 1980b). Similarly, they are abundant in the Late Cretaceous amber deposits (New Jersey and Canada ambers: see respectively [START_REF] Grimaldi | A remarkable deposit of fossiliferous amber from the Upper Cretaceous (Turonian) of New Jersey[END_REF]McKellar & Enel, 2012). The only exception is in Orapa, Botswana [START_REF] Brothers | Diversity of Hymenoptera and other insects in the Late Cretaceous (Turonian) deposits at Orapa, Botswana : a preliminary review[END_REF]. It is possible that the shift in proportions between the two superfamilies' occurred earlier than the Cretaceous (see for instance dating estimates of Spasojevic et al, 2021), although this hypothesis would need additional support as no ichneumonoid fossils are known from this period so far. In addition, these changes in diversity and abundance are not underestimated in the fossil record because the relative abundances of the ichneumonoid and evanioid wasps in the Burmese amber biota show that ichneumonoids were slightly less abundant than evanioids [START_REF] Zhang | Hymenoptera (wasps, bees and ants) in mid-Cretaceous Burmese amber: A review of the fauna[END_REF], while ichneumonoids are clearly more abundant than evanioids in the Cenozoic fossil record. However, data from the latest Cretaceous are too poor to be evaluated due to the lack of any deposits.

Nowadays, Ichneumonoidea are hyper-diversified, hyper-abundant, and widespread with more than 44,000 known species, while Evanioidea are clearly less diversified with only ca. 1,400 species described (data compiled from http://fossilworks.org/ and https://hol.osu.edu; both accessed November 25, 2020).! Numerous hypotheses could be proposed to explain this shift. For instance, a host-related diversification may have contributed to this pattern, with a decrease in host abundance favourable to

Evanioidea and an increase in those favourable to Ichneumonoidea. However, any hypothesis will be difficult to verify. It is also worth mentioning that the decline of evanioids (except the family Evaniidae) occurs in parallel with, and is probably correlated to, the diversification of the xylophilous ichneumonoids. In contrast, the rise of Evaniidae parallels that of the modern Dictyoptera that appeared temporally close to the Jurassic-Cretaceous boundary and diversified during Cretaceous and Cenozoic [START_REF] Basibuyuk | The limits of the family Evaniidae (Insecta: Hymenoptera) and a new genus from Lebanese amber[END_REF].

ONE GENUS & TWO DEPOSITS: A DIRECT EVIDENCE OF THE ISOLATION OF THE WEST BURMA BLOCK

The record of three new othniodellithid wasps from two different Cretaceous Burmese amber provides new clues to investigate the stability and the isolation of the West Burma Block (WBB). In fact, the WBB has experimented a tumultuous geological history with a long migration from Gondwana. During the Early Jurassic, the WBB was located near the Australian block in East Gondwana [START_REF] Seton | Global continental and ocean basin reconstructions since 200 Ma[END_REF][START_REF] Van Hinsbergen | Greater India Basin hypothesis and a two-stage Cenozoic collision between India and Asia[END_REF] and was separated from East Laurasia (Asian region) by the Meso-Tethys Ocean. Numerous studies have investigated the break-up of the WBB + Indian block and they estimated that the break-up from the Gondwana + Australian block occurred between the Late Jurassic and the Early Cretaceous [START_REF] Heine | Reconstructing the lost eastern Tethys ocean basin: convergence history of the SE Asian margin and marine gateways[END_REF][START_REF] Heine | Late Jurassic rifting along the Australian North West Shelf: margin geometry and spreading ridge configuration[END_REF]. A recent study [START_REF] Westerweel | Burma Terrane part of the Trans-Tethyan arc during collision with India according to palaeomagnetic data[END_REF] refined these estimates and agreed with previous works [START_REF] Metcalfe | Allochthonous terrane processes in Southeast Asia[END_REF][START_REF] Metcalfe | Pre-cretaceous evolution of SE Asian terranes[END_REF][START_REF] Scotese | Atlas of Early Cretaceous Paleogeographic Maps, PALEOMAP Atlas for Arc-GIS[END_REF], suggesting that the separation dated back ca. 125 Ma.

They also assumed that the Terrane became an isolated island only after ca. 120 Ma. During the resin formation and production, both deposits (Hkamti and Noije Bum) were isolated geographically and occupied an island position in the Tethys Ocean [START_REF] Westerweel | Burma Terrane part of the Trans-Tethyan arc during collision with India according to palaeomagnetic data[END_REF], during more than 20 Ma [START_REF] Heine | Reconstructing the lost eastern Tethys ocean basin: convergence history of the SE Asian margin and marine gateways[END_REF][START_REF] Seton | Global continental and ocean basin reconstructions since 200 Ma[END_REF][START_REF] Licht | Magmatic history of central Myanmar and implications for the evolution of the Burma Terrane[END_REF].

Interestingly, the geological history of the WBB and the othniodellithid fossil record confirm this isolation. In fact, the †Othniodellithidae are endemic to the Burmese amber biota and representatives of the family are now found in two Cretaceous Burmese deposits of different ages, respectively early Albian and Cenomanian. These records are direct evidence of the stability of the WBB biota during at least 10 Ma, which also supports the hypothesis of a long journey of the isolate plate through the Tethys Ocean from Gondwana to Asia. These new discoveries suggest that the WBB has remained isolated long enough for the development of a unique biota. If a part of Burmese amber biota shares affinities with the South Gondwana (e.g. [START_REF] De | Earliest onychophoran in Amber Reveals Gondwanan Migration Patterns[END_REF][START_REF] Poinar | Burmese amber: evidence of Gondwanan origin and Cretaceous dispersion[END_REF]Jouault and Nel, 2020) or Laurasia fauna (e.g. [START_REF] Gumovsky | Laurasian ancestors and "Gondwanan" descendants of Rotoitidae (Hymenoptera: Chalcidoidea): what a review of Late Cretaceous Baeomorpha revealed[END_REF][START_REF] Martynova | Description of the first sclerogibbid wasp (Hymenoptera: Sclerogibbidae) from Burmese (Myanmar) amber and its phylogenetic significance[END_REF][START_REF] Jouault | Mid-Cretaceous Burmese amber pelecinid wasps (Hymenoptera, Pelecinidae) support the hypothesis of an Asian origin of the family[END_REF], an even larger part is endemic and results from the isolation of WBB for at least 10 to 20 Ma.

CONCLUSION

Our study presents the currently most complete phylogenetic hypothesis of Evanioidea, including most extinct genera, and provides divergence times for all known evanioid families.

Bayesian inferences support the monophyly of the Evanioidea and of the families †Anomopterellidae, †Othniodellithidae, †Andreneliidae, Aulacidae, Gasteruptiidae, and Evaniidae. Praeaulacidae and †Baissidae appear to be paraphyletic. The Bayesian tree also allows the transfer of Vectevania and Hyptiogastrites among Aulacidae. The inferred divergence times of Evanioidea indicate that they have probably originated in the Late Triassic-Early Jurassic, which corroborates previous estimations. The evolutionary history of evanioid wasps seems to be a succession of appearances and extinctions of families, the most recent replacing the oldest in their ecological niches or conquering new ones thanks to clade-specific adaptations (viz. †Othniodellithidae). Additionally, the relative abundance and diversity of the evanioid and ichneumonoid wasps in the deep past seem to follow an opposite pattern. However, further works are necessary to deepen our understanding of the various drivers of evanioid diversification, especially in relation to their competitors and hosts.

  Published work and nomenclatural acts are registered in ZooBank (http://www.zoobank.org/, last a c c e s s : 1 5 M a r c h 2 0 2 1 ) , w i t h t h e f o l l o w i n g L S I D ( r e f e r e n c e ) : urn:lsid:zoobank.org:pub:9E9E6EC8-29E8-488A-AF43-0619A1820A7A SYSTEMATIC PALEONTOLOGY CLASS INSECTA LINNAEUS, 1758 ORDER HYMENOPTERA LINNAEUS, 1758 SUBORDER APOCRITA GERSTAECKER, 1867 SUPERFAMILY EVANIOIDEA LATREILLE, 1802 FAMILY †OTHNIODELLITHIDAE ENGEL & HUANG, 2016 GENUS KERADELLITHA JOUAULT, MARECHAL, WANG & PERRICHOT, gen. nov. urn:lsid:zoobank.org:act:1CFB1DFA-B7F1-4414-8CAD-67515F278530 Type species: Keradellitha basilici Jouault, Maréchal, Wang & Perrichot, sp. nov. Other species included: Keradellitha anubis Jouault, Maréchal, Wang & Perrichot, sp. nov.; Keradellitha kirina Maréchal, Jouault & Perrichot sp. nov.

  Jouault, Maréchal, Wang & Perrichot sp. nov. urn:lsid:zoobank.org:act:D217B456-4EEF-4EF7-87B9-A497EE9BCA66 (Figs 3-4) Holotype: NIGP174739, preserved in a rectangular piece of amber measuring 5 × 5 × 4 mm. Type locality and horizon: Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; upper Albian to lower Cenomanian, mid-Cretaceous.

  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 mid-length to apical segment. Parameres directed backward, keel-shaped, protruding about 0.2 mm. Aedegus not clearly visible. Female unknown. KERADELLITHA KIRINA Maréchal, Jouault & Perrichot sp. nov. urn:lsid:zoobank.org:act:47865DDE-514F-4583-AB69-FF0D02A3BD5D Holotype: IGR.BU-021, preserved in rectangular piece of amber measuring 9 × 6 × 3 mm. Type locality and horizon: Hkamti site, Hkamti District, Sagaing Region, Myanmar; early Albian (ca. 110 Ma), early Cretaceous.

Figure 1 .Figure 2 .

 12 Figure 1. Keradellitha basilici Jouault, Maréchal, Wang & Perrichot gen. et sp. nov., holotype NIG-

Figure 3 .

 3 Figure 3. Xenodellitha anubis Jouault, Maréchal, Wang & Perrichot gen. et sp. nov., holotype NIG-

Figure 4 .

 4 Figure 4. Xenodellitha anubis Jouault, Maréchal, Wang & Perrichot gen. et sp. nov., holotype NIG-

Figure 5 .

 5 Figure 5. Xenodellitha kirina Maréchal, Jouault & Perrichot gen. et sp. nov., holotype IGR.BU-021

Figure 6 .

 6 Figure 6. Xenodellitha kirina Maréchal, Jouault & Perrichot gen. et sp. nov., holotype IGR.BU-021

  apical teeth in Othniodellitha mantichora); the gena shorter than compound eyes (vs. wider than compound eyes in O. mantichora); the fore wing with M+Cu and Cu aligned (vs. non-aligned in O. mantichora), 2cu-a and 2m-cu present (vs. both veins absent in O. mantichora), 2rs-m tubular (vs. non tubular in the other genera), 2A present (vs. absent in O. mantichora and Xenodellitha preta); the hind wing with five or more hamuli (vs. three in Xenodellitha); and legs with tarsal plantulae present (vs. absent in the other genera, but maybe not mentioned). 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 ……………….. Keradellitha basiliciJouault, Maréchal,

	Wang & Perrichot sp. nov.
	2. Fore wing with third and second submarginal cells similar in length, second discal cell about
	twice as long as wide …………………………………………….… Keradellitha kirina Maréchal,
	Jouault & Perrichot sp. nov.
	KEY TO GENERA OF †OTHNIODELLITHIDAE:
	1. Flagellum with 20 flagellomeres; hind wing with three hamuli …… Xenodellitha Engel, 2017
	-Flagellum with more than 20 flagellomeres; hind wing with five or more hamuli …………….… 2
	2. Flagellum with 21 flagellomeres; cephalic horn with apical teeth; fore wing with M+Cu not
	aligned with 1Cu, 2m-cu absent, 1M and 1cu-a not aligned, 2cu-a absent, 2A absent
	………………………………………..……………………….. Othniodellitha Engel & Wang, 2016
	1.	-Flagellum with 22 flagellomeres; cephalic horn without apical teeth; fore wing with M+Cu
		aligned with 1Cu, 2m-cu present, 1M and 1cu-a aligned, 2cu-a present, 2A present
		……………………………… Keradellitha Jouault, Maréchal, Wang & Perrichot gen. nov.
		The three species of Keradellitha gen. nov. can be distinguished mainly by the characters
	given in the key below.

KEY TO SPECIES OF KERADELLITHA gen. nov.:

1.

-Fore wing with third submarginal cell longer than second one, second discal cell conspicuously longer than wide (three times) …………..………………… Keradellitha anubis Jouault, Maréchal, Wang & Perrichot sp. nov. Additionally, K. anubis differs from K. basilici

Jouault, Maréchal, Wang & Perrichot gen. et sp. nov.

Table and Figure captions Table 1 .

 andcaptions1 Diversity and distribution of evanioid fossil genera.

Table 2 .

 2 Comparison of results from maximum parsimony (MP) analysis and Bayesian analyses (BI).

Table 3 .

 3 Hierarchical, and suprageneric classification of Evanioidea.

Table 4 .

 4 Divergence time estimates of Evanioidea and its constitutive families under different models and priors.

Table 2 .

 2 Comparison of results from maximum parsimony (MP) analysis and Bayesian analyses (BI).

	Families/Subfami-	Species	Distribution	Period	References
	lies / Genera
		8	China	Middle Jurassic	Zhang & Rasnitsyn,
					2008; Li et al.
					2013b; Li et al.
					2014b
	Choristopterella Li	1	Kazakhstan	Middle Jurassic	Rasnitsyn, 1975
	et al.
	Synaphopterella Li	1	China	Middle Jurassic	Li et al. 2013a
	et al.
	Baissidae
	Baissa Rasnitsyn	2		Early Cretaceous	Rasnitsyn, 1975,
					1991
		1	Mongolia	Early Cretaceous	Rasnitsyn, 1991
	Electrobaissa Engel 1	USA	Late Cretaceous	Engel, 2013
	Heterobaissa Li et	1	China	Early Cretaceous	Li et al. 2018
	al.
	Humiryssus Lin	5	China	Early Cretaceous	Lin, 1980; Hong &
					Wang, 1990; Zhang
					and Rasnitsyn 2004
		1	United Kingdom	Early Cretaceous	Rasnitsyn et al.,
					1998
	Manlaya Rasnitsyn	3	China	Early Cretaceous	Ren, 1995; Li et al.,
					2019
		8	Mongolia	Early Cretaceous	Rasnitsyn, 1980a;
					1986
		2	Russia	Early Cretaceous	Rasnitsyn, 1980b
		2	Spain	Early Cretaceous	Rasnitsyn &
					Martínez-Delclòs,
					2000
		3	United Kingdom	Early Cretaceous	Rasnitsyn et al.,
					1998
	Mesepipolaea Zhang	2	China	Early Cretaceous	Zhang & Rasnitsyn,
	& Rasnitsyn				2004; Li et al., 2019
	Tillywhimia Rasnit-	2	United Kingdom	Early Cretaceous	Rasnitsyn et al.,
	syn & Jarzembowski				1998

Table 4 .

 4 Divergence time estimates of Evanioidea and its constitutive families under different models and priors.

		MKv_-	MKv_u-	MKvG_-	MKvG_-	MKvG_-	MKv-	MKv-	MKv-	MKv-
		fixed_-	ni_fos-	fixed_-	fixed_-	fixed_-	G_uni_-	G_uni_-	G_uni_-	G_uni_-
		Fossiltip	siltip	diversity	Fossiltip	random	diversity	flatprior	fossiltip	random
		me an	95 % HP D	me an	95 % HP D	me an	95 % HP D	me an	95 % HP D	me an	95 % HP D	me an	95 % HP D	me an	95 % HP D	m ea n	95 % HP D	me an	95 % HP D
	St	20	18	20	19	22	20	19	18	23	20	23	21	20	18	20	18	24	22
	e	2	7-	7	1-	8	6-	8	4-	1	8-	9	5-	3	8-	3	8-	9	6-
	m-		21		22		25		21		25		26		22		22		27
	Ev		8		3		3		4		4		2		0		0		5
	ani
	oi
	de
	a
	Cr	17	16	17	16	19	17	17	15	19	18	20	18	17	16	17	15	21	19
	ow	5	0-	8	1-	7	8-	3	6-	9	0-	6	6-	6	0-	6	9-	4	3-
	n-		19		19		21		18		22		22		19		19		23
	Ev		2		6		9		9		1		7		4		4		8
	ani
	oi
	de
	a
	Pr	19	18	19	19	21	19	19	18	21	19	22	20	20	18	20	18	22	20
	ot	9	5-	6	1-	4	4-	5	1-	6	4-	4	2-	0	5-	1	6-	7	5-
	oe		21		22		23		21		23		24		21		21		24
	va		4		3		7		0		7		6		5		5		9
	ni
	oi
	de
	s
	Pr	19	18	19	19	21	19	19	18	21	19	22	20	20	18	20	18	22	20
	ae	9	5-	6	1-	4	4-	5	1-	6	4-	4	2-	0	5-	1	6-	7	5-
	aul		21		22		23		21		23		24		21		21		24
	aci		4		3		7		0		7		6		5		5		9
	da
	e
	Ot	11	11	11	10	11	11	11	11	11	11	12	11	11	11	11	11	12	11
	hni	6	0-	7	9-	7	0-	6	0-	8	0-	0	1-	7	0-	7	0-	3	3-
	od		12		12		12		12		12		13		12		12		13
	elli		6		7		7		5		8		0		7		7		5
	thi
	da
	e

ACKNOWLEDGEMENTS

This study was conducted during a research internship of CJ and AM as part of their Master thesis (under the supervision of VP). We thank Prof. Alexandr P. Rasnitsyn and an anonymous reviewer for insightful comments on the manuscript. This research was supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB26000000), the Second Tibetan Plateau Scientific Expedition and Research (2019QZKK0706), and National Natural Science Foundation of China (41688103).

and Cenozoic (data from http://fossilworks.org/).


Corentin Jouault
email: jouaultc0@gmail.com

Andre Nel
email: anel@mnhn.fr@orcid.org

A new species of Prosyntexis Sharkey 1990 (Hymenoptera: Sepulcidae) from Crato Formation confirmed by geometric morphometric analysis

Keywords: Symphyta, morphometry, Santana, Cretaceous, Aptian

. The new species can be differentiated from the other Prosyntexis species of the Crato formation by its larger size but also in having the cell 2R1 small, the cell 2M small and short, the cell 3R1 narrow and the vein a directed toward wing apex. We performed a Geometric Morphometric Analysis (GMA) to estimate the variation in the forewing venation of the two species from the Crato Formation and ensure our placement.

Introduction

The Sepulcidae [START_REF] Rasnitsyn | Noviye Mezozoyskiye Pilil'shchiki (Hymenoptera, Symphata) [New Mesozoic sawflies (Hymenoptera, Symphata)]. Yurskie Nasekomiye Karatau [Jurassic Insects of Karatau[END_REF] is an important Mesozoic wasp family with several species ranging in age from the Jurassic [START_REF] Rasnitsyn | Noviye Mezozoyskiye Pilil'shchiki (Hymenoptera, Symphata) [New Mesozoic sawflies (Hymenoptera, Symphata)]. Yurskie Nasekomiye Karatau [Jurassic Insects of Karatau[END_REF] to the Cretaceous (e.g., [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF][START_REF] Darling | Insects from the Santana formation, Lower Cretaceous, of Brazil[END_REF][START_REF] Jattiot | Revision of Prosyntexis from the Lower Cretaceous Crato Formation of Brazil (Hymenoptera: Sepulcidae: Trematothoracinae)[END_REF][START_REF] Kopylov | New Trematothoracinae (Hymenoptera: Sepulcidae) from the Lower Cretaceous of Transbaikalia[END_REF], 2017a, b). The representatives of this fossil family are all described from fossil imprints, challenging observations of numerous key characters due to the incomplete state of preservation of the fossils (see, e.g., [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF][START_REF] Rasnitsyn | New Early Cretaceous hymenopterous insects (Insecta: Hymenoptera) from Sierra del Montsec (Spain)[END_REF][START_REF] Rasnitsyn | Wasps (Insecta: Vespida = Hymenoptera) from the Early Cretaceous of Spain[END_REF][START_REF] Zhang | Discovery of Trematothoracinae (Insecta) in the Upper Jurassic of W. Liaoning, China with a discussion of its phylogeny[END_REF].

Within the family, the Lower Cretaceous genus Prosyntexis Sharkey, 1990 comprises species described from Brazil (e.g., [START_REF] Jattiot | Revision of Prosyntexis from the Lower Cretaceous Crato Formation of Brazil (Hymenoptera: Sepulcidae: Trematothoracinae)[END_REF], Mongolia [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF], Russian Federation [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF] and Spain [START_REF] Rasnitsyn | New Early Cretaceous hymenopterous insects (Insecta: Hymenoptera) from Sierra del Montsec (Spain)[END_REF]. The type species P. gouleti [START_REF] Darling | Insects from the Santana formation, Lower Cretaceous, of Brazil[END_REF] was described from fossil imprints originating from the Crato Formation of Brazil [START_REF] Darling | Insects from the Santana formation, Lower Cretaceous, of Brazil[END_REF]. This fossil locality is known to be one of the largest fossil insect Lagerstätten and yields numerous symphytan specimens resulting in the description of numerous species (e.g., [START_REF] Jattiot | Revision of Prosyntexis from the Lower Cretaceous Crato Formation of Brazil (Hymenoptera: Sepulcidae: Trematothoracinae)[END_REF][START_REF] Freitas | New species of Myrmicium Westwood (Psedosiricidae = Myrmiciidae: Hymenoptera, Insecta) from the Early Cretaceous (Aptian) of the Araripe Basin, Brazil[END_REF][START_REF] Jouault | The first fossil horntail wasp (Hymenoptera: Siricidae) from Lower Cretaceous Crato Formation in Brazil[END_REF]. Recently, Prosyntexis legitima [START_REF] Martins-Neto | A new species of wasp (Symphyta, Sepulcidae) from the Santana Formation (Lower Cretaceous, Northeast Brazil)[END_REF] was described from the same outcrop but was later synonymized under P. gouleti [START_REF] Jattiot | Revision of Prosyntexis from the Lower Cretaceous Crato Formation of Brazil (Hymenoptera: Sepulcidae: Trematothoracinae)[END_REF]. Using Geometric Morphometric Analysis we confirm this synonymization, explore the wing venation variations of P. gouleti, and show that the diversity of the genus Prosyntexis is underestimated in the Crato Formation.

Herein we describe a new species of Prosyntexis from the Crato Formation based on a wellpreserved specimen. We combine our description to a Geometric Morphometric Analysis (GMA) to ensure our placement.

Material and methods

Origin and preparation of the material

The holotype of Prosyntexis sennlaubi sp. nov. derives from the Crato Formation, also called Crato Member of the Santana Formation. This deposit is composed of a limestone outcropping near Nova Olinda in the northeast Brazilian province of Ceará (7.2° S, 39.4° W: paleocoordinates 12.2° S, 10.8° W) (Barling et al., 2015: fig. 1;[START_REF] Ribeiro | Towards an actualistic view of the Crato Konservat-Lagerstätte paleoenvironment: A new hypothesis as an Early Cretaceous (Aptian) equatorial and semi-arid wetland[END_REF] fig. 1). The fossiliferous unit has been dated as late Aptian on the basis of its palynology [START_REF] Pons | Quelques observations sur la palynologie de l!Aptien supérieur et de l!Albien du bassin d!Araripe (N.E. du Brésil)[END_REF][START_REF] Varejão | Mixed siliciclastic-carbonate sedimentation in an evolving epicontinental sea: Aptian record of marginal marine settings in the interior basins of north-eastern Brazil[END_REF]. The fossil is a partly compressed, three-dimensional mineralized replica. The cuticle was replaced by dark brown iron hydroxide (goethite); internally, the insect body is composed largely of calcite and apatite.

The holotype of Prosyntexis sennlaubi sp. nov. was prepared by removing the limestone matrix from around the body with fine blades and needles; the limestone dust was then removed with slight puffs of compressed air. Figures were composed with Adobe Illustrator CC 2019 and Photoshop CC 2019 software. We follow the wing venation of [START_REF] Jouault | The first fossil horntail wasp (Hymenoptera: Siricidae) from Lower Cretaceous Crato Formation in Brazil[END_REF].

Geometric Morphometric Analyses (GMA)

GMA is often used in phenetic studies of relationships but rarely to explore the disparity and variations between species. Herein, we used seven forewings belonging to four specimen of Prosyntexis originating from the Crato Formation (Tab. 1). Based on the criterion for GMA that all specimens must have the same number of landmark points, we are limited by using only a reduced series of figured and non-figured specimens with well-preserved forewing venations. The original images were processed using ImageJ software [START_REF] Schneider | NIH Image to ImageJ: 25 years of image analysis[END_REF] for 16 landmarks of forewings of Prosyntexis, all landmarks representing key points on the wing veins (e.g., relative positions and shapes of the veins, branching points and/ or cells; Fig 1). The xy coordinates from wings were standardized using the 'Procrustes' function implemented in ImageJ. We generated a covariance matrix from the 16 landmarks and analysed the differences responsible for the variations using a principal components analysis. The two principal component axes (PC) were the most informative representing nearly 80% of the variation (Fig. 2A,B). We exported the PC scores and analyzed them using the software PAST v4.05 [START_REF] Hammer | PAST: Paleontological statistics software package for education and data analysis[END_REF]) performing a 'multivariate analysis' on the first two components using the parameter 'Clustering' and 'classical', with the unweighted pair-group method using the arithmetic averages (UPGMA) method and the similarity index set as 'Euclidean'. The created tree shows the phenetic relationships among the studied specimens (Fig. 2C). Other species. Prosyntexis gobiensis [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF] (early Aptian, Bon Tsagaan, Mongolia), Prosyntexis okhotensis [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF] Etymology. The specific epithet honors Markus Sennlaub who kindly allowed the specimen study.

Diagnosis.

Large wasps; cell 2R1 small, about 1.78 times longer than wide; cell 2M small and short, 1.54 times longer than wide; cell 3R1 narrow, 3.57 times longer than wide.

Locality and age. Early Cretaceous, late Aptian or perhaps early Albian, Crato Formation, Brazil, Araripe Basin.

Description. Large wasp with a wing span ca. 27 mm and body length as preserved about 14.42 mm long and about 2.52 mm wide (measure at the level of wing insertion). Surface sculpture of body not apparent. No distinctive color pattern; as preserved, body black-to brown-colored; forewings somewhat infuscate, fully corrugate with pterostigma and vein Sc darker than surrounding veins.

Head rounded, nearly as long as wide (1.89 mm long, 1.95 mm wide with eyes included); eyes occupying most of head lateral surface, ca. 1.10 mm long; antenna inserted anteriorly on head with at least 10 preserved segments; vertex convex; ocelli not preserved.

Thorax (in ventral view) ca. 2.52 mm wide; tergite and sternite not discernible; fore and mid legs partially preserved and apparently slender; hind femora ca. 2.70 mm long, hind tibia ca. 2.95 mm long slightly widening towards apex; hind tarsus at least 2.24 mm long, tarsi not measurable.

Forewing ca. 12.30 mm long, ca. 4.30 mm wide (measure in apical third). Membrane covered with corrugations; both forewings almost complete with costa not perfectly preserved; costal space narrow, slightly enlarged medially (maybe due to preservation); C darkened, not fully visible; R straight; M+Cu and 1-A slightly curved basally; pterostigma broad and sclerotized; first abscissa of Rs (1-Rs) nearly straight, originating after Sc+R mid-length, 0.84 times as long as 1-M; 1-M shorter than Rs+M and 1-Cu veins; 1r-rs inclined and subequal to 1r-m, about twice as long as 2r-rs, cell 2R1 small; 2r-rs straight and directed towards wing apex; 3r-m straight, slightly inclined, and separated from 2r-rs by a length nearly equal to 1.7 of length of 3r-m, posteriorly located after 2M cell; 1m-cu vein nearly straight, shorter than 1-Rs and separating 1M and 2M cells; 1M cell rectangular with parallel sides; 1cu-a straight, located at base of cell 1M, nearly aligned to slightly postfurcal to 1M; 2M cell trapezoidal and elongate, 1.52 times longer than cell 1M, closed by a straight 2m-cu vein slightly inclined toward wing apex; 2m-cu situate in apical third of 1+2Rs cell, at a distance longer than its length from 1m-cu; 1Cu cell elongate, basally narrowing with M+Cu and 1-A veins approaching at base (maybe with a short vein stub emerging from 1-A in its distal part, only visible on right wing), separated from 2Cu cell by 1cu-a vein; 2Cu broadly triangular, elongate, tapering toward wing apex; 1A vein long, anal loop incomplete and not well-marked; 2+3A long, straight meeting 1A apically with a least one crossvein a; anal cells not visible (preservation incomplete or leg obscuring view). Hind wings partially preserved.

Abdomen incomplete with preserved part at least 8.20 mm long and 2.95 mm wide, slightly tapering toward apex, with at least eight segments.

Sex: female due to the presence of a medial carina on the last abdominal segments that received the ovipositor.

Remark.

There is an important difference in the coloration of the forewing veins C and R between P. gouleti and P. sennlaubi sp. nov., viz. the former has both C and R dark while P. sennlaubi (Fig. 4) has C dark and R pale. But in the case of P. gouleti, the original colors are no longer present as the organic matter was replaced by iron oxide.

The forewing length of Prosyntexis sennlaubi sp. nov. is equal to 14.42 mm while the maximal length of the forewing of Prosyntexis gouleti is equal to 11.5 mm (average length near 9.66 mm measured on known specimens).

Discussion

The forewing venation of the fossil specimens described and figured above are nearly identical to those of the known representatives of the genus Prosyntexis (Fig. 6), currently included in the sepulcid subfamily Trematothoracinae. However, the body, the head and the antennae are poorly preserved in Prosyntexis sennlaubi sp. nov., a similar preservation is recorded in numerous specimens of P. gouleti including the one shown here. [START_REF] Darling | Insects from the Santana formation, Lower Cretaceous, of Brazil[END_REF] proposed the following diagnosis for the genus Prosyntexis: 2r-m crossvein of forewing absent; 2R1 cell of forewing wider basally than apically; 1Rs+M of forewing reaching lm-cu crossvein; 3R1 cell closed; 1M cell long and four-sided; cu-a crossvein only slightly postfurcal (Fig. 6). All these character states are found in our new species supporting its attribution to the genus Prosyntexis. However, the character 'Rs+M of forewing reaching lm-cu crossvein' is not recorded in our specimen of P. gouleti since 1Rs+M forks slightly basal of 1m-cu. This state was also mentioned and similar in another specimen of P. gouleti, ensuring our placement (see Jattiot et al., 2011: figs 1-10).

Sharkey (in

Prosyntexis sennlaubi sp. nov. differs from P. gouleti in having longer and broader forewings (ca. 12.30 mm long vs. 9.66 mm long on average in P. gouleti); smaller cell 2R1; thinner cell 3R1; shorter 2M cell and 1A cell closed basally by vein a directed toward wing apex (vs. directed toward wing base). Additionally, the wing venation analysis under GMA indicates that Prosyntexis sennlaubi sp. nov. occupies a morphospace separate from P. gouleti. Prosyntexis sennlaubi sp. nov. differs from Prosyntexis gobiensis [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF] in having the veins 3r-m and 2m-cu not aligned and the cell 2R1 longer than wide (Rasnitsyn, 1993: fig. 28). It differs from Prosyntexis okhotensis [START_REF] Rasnitsyn | Novye taksony sepulek (Vespida: Sepulcidae) [New taxa of Sepulcidae (Mesozoic Vespida from Asia)[END_REF] in having a more elongate cell 1+2Rs and a cell 1M longer than wide (Rasnitsyn, 1993: fig. 27). The forewing venation of Prosyntexis montsecensis Rasnitsyn and Ansorge, 2000 is very incompletely preserved; nevertheless its cell 3R1 is narrower than in Prosyntexis sennlaubi sp. nov., compared to its length (Rasnitsyn & Ansorge, 2000: fig. 1b).

Conclusion

The species of the genus Prosynthexis differ in few characters, mainly in the proportions and shapes of the forewing cells and veins. Nevertheless, we could demonstrate that the use of the phenetic Geometric Morphometric Analysis tools can be efficient to detect a new, 'cryptic' species among these wasps. It is remarkable that so similar species were present in very distant places during the Barremian to Aptian of Mongolia, Spain, and Brazil. During all the Early Cretaceous, South America was still connected to Africa but separated from Spain (an island close to Europe) by the Tethys Ocean, while Mongolia, in Eastern Asia, was more or less connected to North America by a 'Beringian' bridge, while North and South America were separated (Scotese, 2014a, b). It is only during the Latest Jurassic (around 145.5 Ma) that all these regions was better connected, allowing faunal exchanges, suggesting a Latest Jurassic origin and a remarkable morphological stability during ca. 30 Ma for the genus Prosynthexis.

  This published work and new nomenclatural acts are registered in ZooBank with the following LSID (reference): urn:lsid:zoobank.org:pub:C6C329DD-25A4-46A4-96BA-A02036B4D0BE Prosyntexis gouleti Sharkey, 1990 (late Aptian, Sierra do Araripe, Brazil).

  (early Aptian, Bon Tsagaan, Mongolia), Prosyntexis montsecensis Rasnitsyn & Ansorge, 2000 (early Barremian, Montsec, Spain), Prosyntexis sennlaubi sp. nov. (late Aptian, Sierra do Araripe, Brazil). Prosyntexis gouleti Sharkey, 1990 Studied material. Specimen MNHN.F.A43720 (coll. Borschukewitz), deposited in the Laboratoire de Paléontologie, Muséum national d!Histoire naturelle, Paris, France; Specimen SMNS 66301 and specimen SMNS 66698a (imprint) -66698b (counterimprint), deposited in the fossil collection of the State Museum of Natural History Stuttgart, Germany; Specimen 18022 from M. Sennlaub private collection (Figs 3, 6A). Prosyntexis sennlaubi sp. nov. (Figs 4, 5, 6B) urn:lsid:zoobank.org:act:2F40B159-B8A5-47E3-932F-417BEF8E5661 Holotype. Specimen identifier number 18.036 (a well-preserved 3-D fossil in ventral view), Markus Sennlaub will deposit it in the future in the Musée d!Histoire Naturelle et d!Ethnographie, Colmar, France.

FIGURE 1 .

 1 FIGURE 1. Prosyntexis sennlaubi sp. nov., holotype 18.036, forewing with landmarks used in

FIGURE 2 .

 2 FIGURE 2. Results of the GMA analysis. A, Percentage of variance related to the principal

FIGURE 3 .

 3 FIGURE 3. Prosyntexis gouleti Sharkey, 1990 Specimen 18.022. A, Photograph. B, Interpretative

FIGURE 4 .FIGURE 5 .

 45 FIGURE 4. Prosyntexis sennlaubi sp. nov. holotype 18.036, photograph of habitus. Scale bar =

FIGURE 6 .

 6 FIGURE 6. Interpretative line drawing of forewing of Prosyntexis spp. from Crato formation. A, P.

Table 1 .

 1 Material used in the GMA analysis.

Acknowledgements

We thank a lot Prof. A.P. Rasnitsyn, an anonymous referee, and Dr. Neal Evenhuis for their useful remarks on the first version of the paper. We particularly thank Mr. Markus Sennlaub who allowed us to study the holotype specimen.

Species