Corentin Jouault
email: jouaultc0@gmail.com

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

André Nel
email: anel@mnhn.fr

New flat wasps from mid-Cretaceous Burmese amber deposits highlight the bethylid antiquity and paleobiogeography

Keywords: Conceptualization, Data curation, Investigation, Methodology, Visualization, Writing -original draft, Writing -review Hkamti amber, Bethylidae, †Lancepyrinae, paleobiogeography, Abian

New flat wasps from mid-Cretaceous Burmese amber deposits highlight the bethylid antiquity and paleobiogeography (Hymenoptera: Chrysidoidea)

Introduction

The †Lancepyrinae are a small extinct bethylid subfamily, originally described from Barremian Cretaceous Lebanese amber [START_REF] Azevedo | A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position[END_REF]. Members can be easily distinguished from related taxa mainly by having fore wing venation with an unusual combination of closed lanceolate cell 2R1, tubular and well-pigmented vein Rs+M, and angled vein Rs&M [START_REF] Azevedo | A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position[END_REF].

Later, this work was supplemented by the additions of Ortega and [START_REF] Ortega-Blanco | Bethylidae from Early Cretaceous Spanish amber (Hymenoptera: Chrysidoidea)[END_REF] and [START_REF] Engel | A new bethylid wasp in Lebanese Early Cretaceous amber (Hymenoptera: Chrysidoidea), with comments on other Mesozoic taxa[END_REF], with descriptions of further lancepyrine genera and species, challenging the constant presence of a tubular Rs+M vein. †Lancepyrinae were hitherto known only by five species from Lebanese, Spanish, and Taimyr amber [START_REF] Evans | Cretaceous aculeate wasps from Taimyr, Siberia (Hymenoptera)[END_REF][START_REF] Azevedo | A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position[END_REF][START_REF] Ortega-Blanco | Bethylidae from Early Cretaceous Spanish amber (Hymenoptera: Chrysidoidea)[END_REF][START_REF] Engel | A new bethylid wasp in Lebanese Early Cretaceous amber (Hymenoptera: Chrysidoidea), with comments on other Mesozoic taxa[END_REF], but remained unknown from the Burmese amber biota. As for the other bethylid fossil subfamilies, we do not have a record of behavior or clue to understand †Lancepyrinae biology [START_REF] Azevedo | Global guide of the flat wasps (Hymenoptera, Bethylidae)[END_REF].

Recently, two phylogenies have been proposed to investigate the relationships between Bethylidae subfamilies (Jouault et al., 2020a;[START_REF] Colombo | Phylogenetic overview of flat wasps (Hymenoptera, Bethylidae) reveals Elektroepyrinae, a new fossil subfamily[END_REF]. However, if their results do not record †Lancepyrinae at exactly the same position, both recover them as an early diverging lineage among Bethylidae. In Jouault et al. (2020a: fig 6), they fall as a part of separate radiation ( †Lancepyrinae + †Protopristocerinae) sister group of the clade (Mesitiinae + Pristocerinae + Epyrinae + Scleroderminae); while Colombo et al. (2020: fig 1B) recorded them as sister group to the clade (Mesitiinae + †Elektroepyrinae + Pristocerinae + Epyrinae + Scleroderminae + †Protopristocerinae). It is therefore probable that new phylogenies will be necessary to clarify this placement, above all a molecular phylogeny updating the results of [START_REF] Carr | Phylogeny of bethylid wasps (Hymenoptera: Bethylidae) inferred from 28S and 16S rRNA genes[END_REF].

Herein, we describe the first records of †Lancepyrinae from two Cretaceous Burmese amber deposits, and we discuss their placement and the implication on the paleobiogeographic history of the subfamily.

Material and methods

J o u r n a l P r e -p r o o f

The amber piece containing the type of Protopyris myanmarensis gen. et sp. nov. derives from the deposits of Hkamti site (about 80 km southwest of the Angbamo site), Hkamti District, Sagaing Region, Myanmar, see detailed map in [START_REF] Zheng | A Late Cretaceous amber biota from central Myanmar[END_REF] supplementary figure 2) or in Xing and Qiu, (2020: fig. 1). Zircon U-Pb analyses established an early Albian age (109.7 ± 0.4 Ma) for Hkamti amber, based on zircon analyses of clastic deposits at the surface [START_REF] Xing | Zircon U-Pb age constraints on the mid-Cretaceous Hkamti amber biota in northern Myanmar[END_REF]. The paleobiota from this deposit is almost unknown, with only one ostracod and one Diapriidae described to date [START_REF] Xing | A gigantic marine ostracod (Crustacea: Myodocopa) trapped in mid-Cretaceous Burmese amber[END_REF]Jouault et al., 2020b). The amber piece containing the type of Burmapyris azevedoi gen. et sp. nov. derives from the deposits of Noije Bum in the Hukawng Valley (26° 29′ N, 96° 35′ E), Kachin State, northern Myanmar (see detailed map in Grimaldi and Ross, 2017: fig. 2). Radiometric data established an earliest 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 corroborate 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 specimens were examined and photographed using a Leica MZ APO stereomicroscope equipped with a Canon EOS 5D Mark II camera. All images are digitally stacked photomicrographic composites of several individual focal planes, which were obtained using HeliconFocus 6.7. The figures were composed with Adobe Illustrator CC 2019 and Adobe Photoshop CS19 software. The specimens are housed in the amber collection of the Geological Department and Museum (IGR) of the University of Rennes, France. We follow the nomenclature, classification, and terminology for morphological features of the body and wing venation of [START_REF] Azevedo | Global guide of the flat wasps (Hymenoptera, Bethylidae)[END_REF].

Main measurements and abbreviations used are as follows: length of fore wing (LFW); length of head, mandibles excluded (LH); width of head (WH); width of frons (WF); height of eye (HE); length of eye (LE); ocello-ocular line (OOL); width of ocellar triangle (WOT); diameter of anterior ocellus (DAO); vertex-ocular line (VOL). The nomenclature of integument sculpture follows [START_REF] Harris | The glossary of surface sculpturing[END_REF].

J o u r n a l P r e -p r o o f Holotype: Specimen identifier IGR.BU-015 (a nearly complete specimen, with right antenna destroyed, in a rectangular piece of amber measuring 5 × 4 × 1.5 mm).

Etymology. Named after the originating country of the amber piece, Myanmar. The specific epithet is to be treated as an adjective in the genitive case.

Locality and horizon. Hkamti site, Hkamti District, Sagaing Region, Myanmar; lower Albian (ca.

110 Ma), mid-Cretaceous.

Diagnosis. As for the genus.

Discussion

Systematic placement and phylogenetic hypothesis

Following the key to superfamilies proposed in [START_REF] Goulet | Hymenoptera of the world: An identification guide to families[END_REF], our specimens key out in Chrysidoidea because of their pronotum being more than half as long as their mesoscutum, and their metasomal terga 1 and 2 having very weak constriction at their junction dorsally and laterally or with no constriction at all. Following the key to Chrysidoidea families in [START_REF] Goulet | Hymenoptera of the world: An identification guide to families[END_REF], they key out in Bethylidae because of their antenna with 10 or 11 flagellomeres, the presence of wings, their profemora not enlarged or swollen, metasoma with six or seven exposed J o u r n a l P r e -p r o o f terga, pronotum touching tegula, and prognathous heads. Additionally, our specimens could be confused with Scolebythidae but differ from the latter in having a pronotal anterior flange rendering propleura visible in dorsal aspect; the propleura closely situated so that the prostrenum is not large, exposed, and diamond-shaped; and the protrochanter inserted ventrally on procoxa [START_REF] Engel | Cretaceous Scolebythidae and phylogeny of the family (Hymenoptera: Chrysidoidea)[END_REF]. Our specimens also differ from †Chrysobythidae at least in lacking a shallow medial line on head (vs. present in †Chrysobythidae); in having the clypeus more or less triangular (vs.

raised as a convex, somewhat squared lobe), the fore wing with a reduced number of closed cells (vs. six enclosed cells) and the cells 1R1 and 2R1 open [START_REF] Melo | †Chrysobythidae, a new family of chrysidoid wasps from Cretaceous Burmese amber (Hymenoptera, Aculeata)[END_REF].

Following the key to subfamily of [START_REF] Azevedo | Global guide of the flat wasps (Hymenoptera, Bethylidae)[END_REF], Protopyris gen. nov. and Burmapyris gen. nov. key out in †Lancepyrinae because of their developed wings; the fore wings with

Rs+M vein tubular and straight, at least as stub, with at most six closed cells; the clypeus not projecting backward onto frons; and the tarsal claws slightly curved. Following the key to genera of †Lancepyrinae of [START_REF] Azevedo | Global guide of the flat wasps (Hymenoptera, Bethylidae)[END_REF], Protopyris gen. nov. keys out near Archaepyris Evans, 1973 because of the fore wing with Rs+M vein conspicuous and tubular, and 2R1 cell opened.

However, Protopyris gen. nov. differs from Archaepyris in having an elongate fore wing, an elongate pterostigma, 1M cell conspicuous and closed, 2Cu cell virtually delimited by spectral veins, and M and cu-a veins aligned. Unfortunately, [START_REF] Evans | Cretaceous aculeate wasps from Taimyr, Siberia (Hymenoptera)[END_REF] did not provide extensive illustrations as for the other species he described. Thus, it is difficult to make a full comparison with our specimen. Following the key to genera of †Lancepyrinae of [START_REF] Azevedo | Global guide of the flat wasps (Hymenoptera, Bethylidae)[END_REF], Burmapyris gen. nov. also keys out near Lancepyris because of the fore wing with Rs+M vein conspicuous and tubular and 2R1 opened, but it differs from the latter in having a fore wing with a long and wide pterostig- J o u r n a l P r e -p r o o f [START_REF] Rasnitsyn | The origin and evolution of Lower Hymenoptera[END_REF][START_REF] Rasnitsyn | Origin and evolution of Hymenoptera][END_REF], proposed a general trend among various hymenopteran lineages consisting in the miniaturization, reduction of venation, enlargement of the pterostigma, and reduction of number of antennal flagellomeres. The reduction of wing venation has been recently investigated in two bethylid subfamilies (viz. †Holopsenellinae and Scleroderminae) by Jouault et al. (2020a), in accordance with Rasnitsyn's observations. Thereby, by comparing the age and the wing venation of the new species described herein and previously described genera, it is possible to infer some phylogenetic placement of Protopyris gen. nov. and Burmapyris gen. nov. (Fig. 5) and to propose a hypothetic phylogeny of the †Lancepyrinae. Lancepyris and Protopyris display a relatively more complete and complex wing venation than other representatives, with numerous closed cells.

Thereby, it is likely that they diverged earlier than other lancepyrine wasps (Fig. 2C) (Azevedo et al., 2018: fig 92D). This hypothesis is also supported by their age, in fact, Lancepyris is the oldest record of the subfamily, which would be congruent with a relatively 'basal' position in the †Lancepyrinae. Proposing relationships between the other taxa is more complicated since the differences in venation are reduced. However, Zophepyris and Burmapyris share a tubular Rs+M vein (Fig. 4D) (Azevedo et al., 2018: fig 92F), not recorded in other 'advanced' genera (viz. Liztor, Archaepyris, Cretepyris), suggesting that they could be closely related. Liztor, Archaepyris, and Cretepyris all have closed R and 1Cu cells (Azevedo et al., 2018: figs 92A,B,C,E), but only Liztor has a reduced 2r-rs&Rs vein while Archaepyris and Cretepyris display an elongated and tubular 2r-rs&Rs vein. Thereby, we assume that these two genera could be closely related to each other than to

Liztor, which could be the sister group of (Archaepyris + Cretepyris) since the rest of their wing venations is similar.

Paleobiogeographic implications

Correlation between paleogeography and fossil record often provided interesting hypotheses on taxa antiquity. The Burma Terrane, where the amber was initially produced, displays an interesting geological and paleobiogeographic history allowing consistent hypothesis on origin and antiqui-J o u r n a l P r e -p r o o f ty of taxa composing the Burmese amber Biota. In fact, during the Early Jurassic, the Terrane was located near the Australian block in East Gondwana [START_REF] Van Hinsbergen | Greater India Basin hypothesis and a two-stage Cenozoic collision between India and Asia[END_REF][START_REF] Seton | Global continental and ocean basin reconstructions since 200 Ma[END_REF] and separated from East Laurasia (Asian region) by the Mesotethys Ocean. In addition, the extant Pacific islands (Malaysia, Papua, Borneo, etc.) did not yet exist [START_REF] J O U R N A L P R E -P R O O F Neall | The age and origin of the Pacific islands: a geological overview[END_REF], making it almost impossible to transfer wildlife from East Laurasia to the East Gondwana (Australia and Burmese Terranes). Following this period, the Burmese Terrane began to break away from the Australian block to go north (as for the Indian block) to the Asian region 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]. The Burma Terrane is supposed to have been separated from the Gondwana after 125 Ma ago, but close to its margin ca. 120 Ma ago [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 ArcGIS[END_REF][START_REF] Westerweel | Burma Terrane part of the Trans-Tethyan arc during colli-J o u r n a l P r e -p r o o f sion with India according to palaeomagnetic data[END_REF]. Later, it was isolated geographically and occupied an island position in the Tethys Ocean between Asia and the Indian block during the mid-Cretaceous [START_REF] Westerweel | Burma Terrane part of the Trans-Tethyan arc during colli-J o u r n a l P r e -p r o o f sion with India according to palaeomagnetic data[END_REF] for 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]. It is very likely that the stem group of the Bethylidae was already present on the Burma Terrane before its complete separation from Gondwana, ca. 125-120 Ma ago, since we already have fossil record of the family in Lebanese amber during the Barremian (Fig. 6) [START_REF] Azevedo | A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position[END_REF]. The record of lancepyrine wasps in Lebanese and Spanish amber (in southern Laurasia), respectively located in the central part of the southern Tethyan margin and the western Tethyan margin (Peris et al., 2016: fig. 1), suggests that the †Lancepyrinae were already diversified along the western Tethys. However, the record of Protopyris gen. nov. and Burmapyris gen. nov. on the Burma Terrane suggests that the †Lancepyrinae had also conquered the eastern Gondwana before the break-up of the Burma Terrane. Cretaceous [START_REF] Azevedo | A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position[END_REF][START_REF] Ortega-Blanco | Bethylidae from Early Cretaceous Spanish amber (Hymenoptera: Chrysidoidea)[END_REF][START_REF] Engel | A new bethylid wasp in Lebanese Early Cretaceous amber (Hymenoptera: Chrysidoidea), with comments on other Mesozoic taxa[END_REF]. The record of Archaepyris in Taimyr amber (Late Cretaceous) [START_REF] Evans | Cretaceous aculeate wasps from Taimyr, Siberia (Hymenoptera)[END_REF] may also indicate that the J o u r n a l P r e -p r o o f †Lancepyrinae already colonized the eastern Laurasia during the mid-Cretaceous. According to this peculiar geological history, it is very likely that the †Lancepyrinae and, therefore the Bethylidae, arose before the break-up of the Burma Terrane from South-East Gondwana, indicating at least an Early Cretaceous (or even Late Jurassic) origin for the family.

Conclusion

Both new genera described herein are the first †Lancepyrinae described from Burmese amber and J o u r n a l P r e -p r o o f

  ma, a tubular Rs+M, 1M and 2Cu cells closed by conspicuous spectral veins, M and cu-a veins aligned, and R1 poststigmal present. Both new genera differ from each other in having the body respectively depressed vs. weakly depressed, and different fore wing venations (see. Figs 1,2C,3,4D).

  Protopyris myanmarensis gen. et sp. nov. is the first Bethylidae described from the lower Albian Hkamti amber. These discoveries highlight the underestimated diversity of bethylid wasps in Burmese biota. The peculiar palaeogeographic history of the Burma Terrane provides evidence for the antiquity of the †Lancepyrinae and, therefore, of the entire Bethylidae family. It is likely that the Bethylidae arose during the earliest Cretaceous, before the Gondwana breakup, and maybe during the Late Jurassic. It stands to reason that a morphological-and molecular-based phylogeny with time divergence estimates has becomes a necessity for our understanding of this family.

  Figure captions.

Figure 1 .

 1 Figure 1. Protopyris myanmarensis sp. nov. Jouault & Nel, holotype IGR.BU-015. Photographs of

Figure 2 .

 2 Figure 2. Protopyris myanmarensis sp. nov. Jouault & Nel, holotype IGR.BU-015. A: Mesosoma.

Figure 3 .

 3 Figure 3. Burmapyris azevedoi sp. nov. Jouault, Perrichot & Nel, holotype IGR.BU-016. Photo-

Figure 4 .

 4 Figure 4. Burmapyris azevedoi sp. nov. Jouault, Perrichot & Nel, holotype IGR.BU-016. A: Head

Figure 5 .

 5 Figure 5. Hypothetical phylogeny of Lancepyrinae with affixed geological time scale.

Figure 6 .

 6 Figure 6. Global paleogeographic reconstruction for the mid-Cretaceous (100 Ma) period, showing

  Etymology. The specific epithet is a patronym honoring Dr. Celso O. Azevedo, a world authority in Bethylidae taxonomy. The specific epithet is to be treat as a noun in a genitive case. crotrichiae and bordered with small microtrichiae, with apical margin gently rounded; anterior border not angularly incurved anterior to pterostigma; veins C, Sc+R, A, R1, Rs, Rs+M, 2r-rs&Rs, M+Cu, M, cu-a and cells C, R, 1Cu, and 2R1 present; M and cu-a veins aligned; cell 2Cu and 1M

	4. Rs+M long, continuing after 1M cell, 1M cell not fully enclosed by tubular
	Burmapyris azevedoi sp. nov. Jouault, Perrichot & Nel veins ……………………… Burmapyris Jouault, Perrichot & Nel gen. nov. (Kachin amber, My-
	urn:lsid:zoobank.org:act:9EE1F4FE-9B6C-43AB-8C2A-823D4A48F0E1 anmar)
	cu-a, Cu and cells C, R, 1Cu, 1M and 2R1 present; M and cu-a veins aligned; cell 2Cu open, virtu-Holotype: Specimen identifier IGR.BU-016 (complete female, in a rectangular piece of amber open, virtually enclosed by spectral vein; cell 2R1 distally opened; Rs+M long and prolonged as a -Rs+M short, only cancelling 1M cell, 1M cell fully enclosed by tubular
	ally enclosed by spectral vein; cell 2R1 distally opened; Rs+M prolonged as a spectral vein origi-measuring 5 × 4 × 2 mm). spectral vein originating from 1M after pterostigma apex; pre-and post-stigmal abscissae of R1 veins ………………………….………. Protopyris Jouault & Nel gen. nov. (Hkamti amber, My-
	nating from 1M at level of pterostigma base; pterostigma long and narrow; prestigmal abscissa of conspicuous. Hind wing with only veins C and Sc+R, and 4 hamuli. anmar)
	R1 conspicuous. Hind wing with only veins C and Sc+R, hamuli uncountable. Metasoma with short petiole; apex interiorized, but six tergites visible, T1 reduced, T2 enlarged, all
	Metasoma with short petiole; six tergites present, T1 reduced, T2 enlarged, all tergites convex, part-Locality and horizon. Noije Bum Hill, Hukawng Valley, Kachin State, Myanmar; upper Albian to tergites convex, partly covering sternites; sting not preserved; length of clearly visible tergites: T1 5. Fore wing with cu-a vein postfurcal with M vein …………………… Zophepyris Engel et al.,
	ly covering sternites; sting not preserved (apical part of metasoma destroyed); length of clearly visi-ble tergites: T1 0.26 mm, T2 0.15 mm, T3 ca. 0.15 mm, more distal tergites impossible to measure because of bent metasoma. lower Cenomanian, mid-Cretaceous. 0.43 mm, T2 0.27 mm, T3 0.22 mm, T4 ca. 0.12 mm, other tergites not measurable since interior-ized. ber, Lebanon) 2016 (Peñacerrada amber, Spain) -Fore wing with cu-a vein aligned or antefurcal with M vein ……………………..……………… 6 6. Fore wing with 2r-rs&Rs vein arising on distal half of pterostig-ma ……………………………... ………………………… .Cretepyris Ortega-Blanco & Engel, 2013 (Peñacerrada amber, Spain) -Fore wing with 2r-rs&Rs vein arising on basal half of pterostig-ma ……………………………….……………………………..… Liztor Ortega-Blanco & Engel, J o u r n a l P r e -p r o o f 2013 (Peñacerrada amber, Spain)
	-Fore wing with 2R1 cell opened …………………………………………………….……….. 3
	3. Fore wing with a short pterostigma and Rs+M vein short (not enclosing 1M cell or reaching apex
	notauli conspicuous; fore wing venation developed, with cells C, R, 1Cu, and 2R1 delimited by Mesosoma with dorsum smooth, not roughly sculptured; femora slightly thickened, tibia thin, tarsal of pterostigma) ………..…………………….… Archaepyris Evan, 1973 (Taimyr amber, Russia)
	tubular veins; M and cu-a veins aligned; pre-and post-stigmal R1 abscissae present; 1M and 2Cu claws simple; tibial spur formula 1-2-2; arolium present; notauli well-marked, slightly posteriorly -Fore wing with an elongate pterostigma and Rs+M long (enclosing 1M cell or reaching apex of
	not clearly delimited; 2R1 opened at apex; pterostigma short, wide in apical one-third of length; convergent; parapsidal furrows weak; metanotum developed medially and overlapping mesoscutel-pterostigma) …………………………………………………………………….………..……… 4

Description. Sex unknown; body ca. 2.10 mm long, depressed, almost smooth and not strongly foveolate; wings clear, setose, LFW 1.25 mm. LH 0.50 mm; WH 0.53 mm; WF 0.31 mm; LE 0.25 mm; OOL 0.15 mm; WOT ca. 0.12 mm; DAO 0.03 mm; VOL 0.11 mm. Head sub-quadrate, not pubescent, with small dorsal median depression, finely punctate but not sculptured; mandible projecting forward, ca. 0.12 mm long, dentate, apparently with four equal, sharp, small teeth along masticatory margin and one longer apical tooth; only two maxillary palpomeres visible (but certainly more are present); flagellomeres with cylindrical cross-section; measurable antennomere lengths (in mm) scape 0.11 mm, pedicel 0.07 mm, F1 ca. 0.03 mm, F2 ca. 0.03 mm, F3 0.05 mm, F4 to F10 sub-equal to equal ca. 0.06 mm (all flagellomeres compressed due to preservation, thereby useful width measurements not provided); occipital carina slightly marked; posterior head margin nearly straight with small median concavity; compound eye without microsetae, closer to mandibular insertion than posterior to head margin; ocellar triangle nearly equilateral, with two posterior ocelli slightly more distant from each other than from anterior.

Mesosoma with dorsum smooth, not roughly sculptured; femora slightly thickened, tibia thin, tarsal claws simple; tibial spur formula 1-2-2; arolium present; notauli well-marked, posteriorly convergent; parapsidal furrows weak; metanotum developed medially and overlapping mesoscutellum posteriorly; metapectal-propodeal complex rectangular, not foveolate, with marginal carinae and no J o u r n a l P r e -p r o o f posterior spines; posterior margin of propodeum straight. Fore wing covered with microtrichiae and bordered with small microtrichiae; elongated with apical margin gently rounded; anterior border not angularly incurved anterior to pterostigma; veins C, Sc+R, A, R1, Rs, Rs+M, 2r-rs&Rs, M+Cu, M, Genus Burmapyris gen. nov. Jouault, Perrichot & Nel Figs 3-4 urn:lsid:zoobank.org:act:A627B4D5-35BB-48F5-B198-A2721DDB01D1 Type species. Burmapyris azevedoi Jouault, Perrichot & Nel sp. nov.

Etymology. The generic epithet is a combination of 'Burma' referring to Burmese amber and the generic name Epyris, commonly used as a suffix for many bethylid genera. Gender masculine.

Diagnosis. Female (sting visible) with body weakly depressed; mandibles heavily dentate with five(?) sharp teeth along masticatory margin, with apical tooth longer and stouter; clypeus triangular and slightly projecting; antennae stout, with 11 flagellomeres; scape nearly twice as long as pedicel; hind wing with four hamuli; propodeum rectangular and not spinose; legs with slightly enlarged femora.

J o u r n a l P r e -p r o o f

Diagnosis. As for the genus.

Description. Female (sting slightly visible); body ca. 2.5 mm long, almost smooth and not strongly foveolate; wings clear and setose, LFW at least 1.50 mm (bent). LH 0.57 mm; WH 0.53 mm; WF 0.31 mm; HE 0.18 mm; LE ca. 0.25 mm; OOL 0.15 mm; WOT ca. 0.15 mm; DAO 0.05 mm; VOL ca. 0.12 mm. Head slightly longer than wide, not pubescent, without dorsal median depression, sculpture apparently absent; mandible bent downward and slightly curved backwards, ca. 0.18 mm long; only four maxillary palpomeres visible (but certainly more numerous); flagellomeres with cylindrical cross-section; measurable antennomere lengths (in mm): scape ca. 0.15 mm, pedicel 0.08 mm, F1 to F7 ca. 0.12 mm, F8 to F10 ca. 0.11 mm, F11 0.18 mm; occipital carina conspicuous; posterior head margin curved; compound eye without microsetae, closer to mandibular insertion than to posterior head margin; ocellar triangle nearly equilateral, situated posteriorly on head, with two posterior ocelli slightly more distant from each other than from anterior. lum posteriorly; metapectal-propodeal complex rectangular, not foveolate, with marginal carinae and no posterior spines; posterior margin of propodeum straight. Fore wing covered with mi-J o u r n a l P r e -p r o o f Key to genera of †Lancepyrinae (modified from Azevedo et al., 2018): 1. Fore wing with Rs+M vein conspicuous and tubular ………………………………………… 2 -Fore wing with Rs+M vein apparently absent, weak or spectral …………………….………… 5 2. Fore wing with 2R1 cell closed …………..…. Lancepyris Azevedo & Azar, 2012 (Ain Dara am-

  Ezcurra and Agnolin (2012: fig. 2) proposed the paleogeographic Eurogondwana model, according to which Gondwanan lineages could colonize Laurasia and North America via an Apulia microplate from 145 to 99 Ma, prior to continental drift. This scenario is in accordance with the presence of other †Lancepyrinae in Spain and Lebanon during the Early

Acknowledgment

We sincerely thank Pr. Alexandr. P. Rasnitsyn, an anonymous referee, and editor Dr. Eduardo Koutsoukos for their very helpful remarks on the first version of the paper.

Declaration of interests

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

J o u r n a l P r e -p r o o f