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Fossils are more and more used in phylogenetic evolutionary studies either for clade calibration, or as terminals in a dataset including morphological characters. The strength of these methodological advances relies however on the quality and completeness of the fossil record. For crickets (Insecta, Orthoptera, Gryllidea), few ancient (pre-Cenozoic) well-preserved fossils are known, except for isolated wings often classified in purely fossil groups and a few fossils found in Cretaceous amber. Here, we present two remarkable fossils from mid-Cretaceous amber of France, that were imaged using X-ray synchrotron microtomography and exhibit an exquisite preservation allowing description with a precision similar to that of extant taxa. Palaeonemobius occidentalis Laurent and Desutter-Grandcolas, gen. nov., sp. nov. and Picogryllus carentonensis Josse and Desutter-Grandcolas, gen. nov., sp. nov. are the oldest representatives of the Nemobiinae and Podoscirtinae subfamilies of the Trigonidiidae and Oecanthidae families respectively. P. carentonensis Josse and Desutter-Grandcolas, gen. nov., sp. nov. is also the smallest adult male with a full stridulatory apparatus ever documented in crickets (body length 3.3 mm), and the first taxon of the cricket clade for which male genitalia can be partly described. We discuss the significance of Cretaceous fossils of crickets for future evolutionary studies of this clade.

The Lancepyrinae are an extinct subfamily of Bethylidae known exclusively from Cretaceous amber deposits of Lebanon, Spain, Taimyr and Myanmar. In this study, we describe and illustrate four new genera and five new species of lancepyrine wasps from the Albian of Hkamti and late Albian-early Cenomanian of Kachin (Myanmar): Azepyris delamarreigen. et sp. nov., Burmapyris ohmkuhnleisp. nov., Gwesped groehnigen. et sp. nov., Paralanceis chotardigen. et sp. nov. and Yunbayin rosseigen. et sp. nov. These taxa not only highlight the taxonomic diversity of the Lancepyrinae during the mid-Cretaceous but they also reveal the morphological disparity of the subfamily. To establish the phylogenetic relationships of these fossils and to check the monophyly of the Lancepyrinae, we add them to a pre-existing morphological matrix and perform a cladistic analysis. We retrieve the subfamily as poorly supported yet monophyletic, with the newly described taxa deeply nested in it. A key to the genera of Lancepyrinae is provided. Finally, we erect the subfamily Cretabythinaesubfam. nov. for the genera Cretabythus Evans, 1973, Holopsenelliscus Engel, 2019 and MegalopsenellaJouault et al., 2020 as no taxonomic treatment has been provided for these taxa after the transfer of HolopsenellaEngel et al., 2016 as Aculeataincertae sedis.

Miocene Dryinidae are relatively well-known compared to other Chrysidoidea families. But most taxa (16 out of 17 described species) derive from the amber mines of the Dominican Republic. Here, a new fossil species of Dryinidae, Thaumatodryinus cotyi sp. nov., is described and figured from the early to middle Miocene of Totolapa (Mexico), and compared to more or less contemporaneous Dominican species. Thaumatodryinus cotyi sp. nov is noteworthy characterized by the testaceous body, the tridentate mandibles, the clypeal anterior margin outcurved, the pronotum striate rugose, the mesopleuron and propodeum areolate rugose, and the propodeum shorter than high. We also propose an up-to-date key to the subfamilies of Dryinidae and summarize the fossil fauna from Totolapa, a deposit hitherto less studied than the coeval amber from Simojovel.

† Desyopone hereon sp. nov. Boudinot and Perrichot (Figures 1–5) ZooBank LSID: urn: lsid:zoobank.org:act: 9E345965-6AA1-42D8-A468-187BABBB38D2. Etymology. The specific epithet gratefully recognizes the Helmholtz-Zentrum Hereon, the research center which funds and operates the Imaging Beamline (P05) at DESY, thus making the present work possible. Holotype. Male (m), MAIG 6016 (Figures 1B, 2A,B, 3 and 4), deposited in the Museum of Amber Inclusions at University of Gda´nsk, Poland. Paratypes. 12 males (m). Synincluded with holotype in amber piece MAIG 6016 (Figures 1A,C,D, 2C,D and 5). Type locality. Exact locality unknown in the Bashilo river gorge near Weldiya, Semien Wollo Zone, Amhara Region, Ethiopia. Type horizon. A fine siltstone/mudstone of Early Miocene age (16–23 Ma). Material. Holotype and paratypes 1–4 are nearly complete, preserved without apparent distortion but with integument entirely covered by a white, opaque (bacterial?) coat for paratypes 1, 2, 4 (Figures 1A,C and 2A). Paratypes 5–12 mostly complete but variously preserved, more or less distorted by apparent dehydration (e.g., Figure 1D). Diagnosis. † Desyopone hereon is uniquely identifiable among all Ponerinae by the distinctly elongate petiolar peduncle and the enlarged, lobate mandibles. Description. Measurements: Holotype (paratypes) (taken from paratypes 1,2,4)—BL 3.11 (3.00–3.50), HW 0.57 (0.50–0.60), HWE 0.70 (0.66–0.70), SL 0.12 (0.10–0.12), EL 0.23, OLL 0.08 (0.08), OIL 0.16 (0.16), WL 1.25 (1.25), ML 0.52 (0.50–0.56), MW 0.49, FWL (2.75), PH 0.26 (0.30), PL ~0.40 (0.40), GL 1.20 (1.00–1.50), GW 0.62 (0.65–0.80), HWI 81 (76–86), SI 21 (20), OCI 50 (50), PI 154 (133). Head (Figures 2–4): Head capsule ovoid-elliptical in lateral view; in full-face view, posterior head margin broadly and evenly convex to compound eyes; oral region of head, i.e., the malar areas, clypeus, and mouthparts, narrower than distance between compound eyes; postgenal bridge short, about 2/5 the length of the head in full-face view as measured from the postocciput to the hypostoma; malar areas distinctly developed; clypeus medially bulging, laterally depressed. Compound eyes situated almost entirely in anterior half of head; eyes subspherical and relatively small, with their length being about 1/3 head length. Ocelli distant from compound eyes, with the lateral ocelli separated from the compound eyes by slightly more than one compound eye length. Antennal toruli located at about head midlength in full-face view, distinctly posterad the posterior clypeal margin. Antennal scapes short, just barely longer than wide, with their length distinctly Mesosoma (Figures 2 and 3): Pronotum short and simple but with distinct muscular convexity as seen in lateral and dorsal views; posterad the anteromedian pronotal lobe (“nuchal lobe”), pronotum in the form of a simple arch, without distinct dorsal and ventral surfaces. Propleurae widely emarginate posteromedially, together forming a broad arch for the prosternum. Prosternum with basisternum apparently arcuate anteriorly; prosternal process developed. Mesoscutum somewhat narrow, with the anteroposterior length slightly greater than the lateromedial width. Notauli developed, and Y-shaped; meeting in the posterior half of the mesoscutum. Parapsides developed, although indistinct. Scutoscutellar sulcus broad, with at least five cross-costae. Mesoscutellum simple, convex, longer than broad in full-face view. Oblique mesopleural sulcus developed. Spiracular lobe (ventrad wing insertions) absent. Mesopleural area divided into dorsal and ventral regions; both regions dorsoventrally taller than anteroposteriorly long. Mesosternal and metasternal regions without processes. Propodeum convex, without armature or distinct sculpturation. Propodeal lobes developed, weak. Legs: Mesotibiae with no spur, metatibiae with a single, pectinate spur. Pretarsal claws narrow and simple, without additional teeth. Arolia well-developed but not very large. Fore wings (Figures 2C,D and 5): Costal vein (C) present, complete. Rsf1 diverging from Sc+ R +Rs well proximad pterostigma, with Sc+ R abscissa about 1/3 pterostigma length. Pterostigma well-developed, long, and narrow, with its length> 5 × its width. Rf distad pterostigma tubular. M+Cu tubular. Mf1 diverging from M+Cu at or slightly distad crossvein cu-a; this abscissa very weakly curved and meeting Rf1 at a distinct oblique angle. Rs+M tubular. Rsf2–3 diverging from Rs+M proximad 1m-cu. Crossvein 1r-rs absent. Crossvein 2r-rs anterior juncture at 2/3 length of pterostigma; this crossvein meeting Rsf proximad crossvein 2rs-m, which is tubular (paratype 4 has 2rs-m duplicated on the right wing, see Figure 2D). Rsf4+ tubular, meeting Rf distally). Mf2 (=abscissa between Rs+M and 1m-cu) short, shorter than 2r-rs and 2rs-m. Mf3 similar in length to but distinctly shorter than Rsf2–3. Mf4+ tubular proximally, becoming nebulous distally, with this occurring at a distance that is about 3 × the length of 2rs-m. Cuf3 (=abscissa of Cu after 1m-cu) joined to 1A posteriorly; 1A is tubular for its entire length. Submarginal cells 1 and 2 similar in size and shape, but with 2 distinctly smaller than 1. Marginal cell 1 long and narrow, with a length that is about 1.5 × pterostigma length. Discal cell 1 subrhomboidal, its length slightly less than 2 × its anteroposterior width. Subdiscal cell closed. Hind wings (Figures 2C and 5): Wing with eight distal hamuli. Jugal lobe absent. C not distinctly developed. R splitting from Sc+ R +Rs distad crossvein 1rs-m; Rf incomplete, not reaching anterior wing margin. Rsf tubular for a distance that is about 2 × the length of Sc+ R +Rs distad 1rs-m. M+Cu splitting well distad crossvein cu-a. Mf1 meeting rs-m at a broad, oblique angle; Mf developed as a stud distad this juncture. Cuf developed distad the split of M+Cu, but exact condition uncertain. Crossvein cu-a long, slightly longer than the length of Mf1; this crossvein situated proximad split of M+Cu by about twice its length. Anal vein (A) tubular past its juncture with cu-a. Anal cell relatively long; M+Cu proximad cu-a distinctly longer than M+Cu distad cu-a. Metasoma (Figures 2B and 3A,D,E): Petiole nodiform and distinctly pedunculate, albeit without a marked inflection between its anterior portion and the anterior surface of the petiolar node; peduncle about 2/5 petiole length; petiolar node height about 3/5 entire petiole length; node broad and convex; posterior collar well-developed; tergosternal fusion absent, laterotergites present; sternum low and very weakly sinuate in lateral view, without an anteroventral (=subpetiolar) process, nor with a posterior process; posterior sternal margin distinctly notched. Helcium infraaxial (=below abdominal segment III midheight); helcial tergite broad and overlapping sternite laterally. Prora not distinctly developed. Gastral segments homonomous in appearance and gradually decreasing in length from abdominal segment III to VIII. Abdominal segment IV without cinctus (=constriction). Abdominal spiracles IV–VIII obscured by preceding tergites. Abdominal tergum VIII apparently simple. Abdominal sternum IX lobate and somewhat narrow, with a length that is about twice its width. Cerci (=pygostyles) developed. Genitalia (Figures 2B and 3A): Cupula present (only visible in µ- CT cross-sections). Gonopods longer than tall. Gonostyli broad proximally, indistinctly set off from gonocoxa, and lobate in appearance, being narrowly rounded apically. Lateropenites (=digiti) thickened apically, downcurved. Penites (=penisvalvae) apparently longer than tall, curving strongly to their narrowly lobate apices. Setation: Body with two primary hair classes: (1) short pubescence, which covers the head and all segments of the antennae, mesosoma, legs, and metasoma; (2) long hairs, which are sparse on all body regions, but are denser on the metasoma., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on pages 9-13, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409

Tribe Ponerini Lepeletier de Saint-Fargeau, 1835. Type genus. Ponera Latreille, 1804. Male diagnosis. In addition to the ponerine plesiomorphies, male Ponerini are distinguished by the following: (1) mandibles vestigial, with an enlarged mandalus, and being variably lobate, spatulate, spiniform, or nub-like (no exceptions known) [synapomorphy, homoplastic among Formicidae]; (2) face between antennal toruli not distinctly raised, thus toruli directed more-or-less dorsally; if the intertorular region is raised, then this region is grooved in appearance due to impression of the supraclypeal area (=frontal triangle) (Plectroctena and Psalidomyrmex with medial torular arches raised, but not face; Hagensia, Megaponera, Ophthalmopone, Simopelta, some Euponera, and some Odontomachus with intertorular region raised) [plesiomorphy]; (3) antennal toruli usually distant from posterior clypeal margin (some Brachyponera, many Leptogenys, Megaponera, and Ophthalmopone have toruli that are close to the clypeal margin) [plesiomorphy]; (4) meso- and metatibiae with two, one, or no spurs; (5) jugal lobe present or absent; (6) helcium distinctly infraaxial, i.e., situated well below the midheight of abdominal segment III (Simopelta is an exception due to softening and reduction in size of the metasoma) [synapomorphy, homoplastic among Formicidae]; (7) abdominal segment IV with or without cinctus; (8) cuticle usually not pruinose, being shiny and variably sculptured (Belonopelta, Hagensia, Megaponera, and Ophthalmopone are exceptions) [plesiomorphy]. Remarks. Definitive infraaxiality in males, i.e., with abdominal tergum III rising high above the petiole, is a strong diagnostic condition for Ponerinae, as this is an infrequent apomorphic condition at the subfamily level. It also occurs in Dolichoderinae and Formicinae, and to some extent in Myrmeciinae and various Myrmicinae. Discothyrea (Proceratiinae) may approach infraaxiality, but the third abdominal tergum is low., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on page 8, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409

Genus † Desyopone gen. nov. Boudinot and Perrichot Type species. † Desyopone hereon sp. nov., by present designation monotypy. ZooBank LSID: urn:lsid:zoobank.org:act: 7228E671-DF5E-44D1-ADE6-62FB6DB34220. Etymology. The genus name is a portmanteau of the traditional ponerine generic suffix, “-pone”, and the acronym for the Deutsches Elektronen-Synchrotron (DESY), whose storage ring and radiation beamline facilities were used to generate the phenomic data that were crucial for the correct identification of the new taxon. Diagnosis. † Desyopone has plesiomorphies 1–4 of Ponerinae and is identifiable as Ponerini at minimum due to the vestigial mandibles and infraaxial helcium. † Desyopone and Cryptopone are uniquely identified among all Ponerinae by: (1) subpetiolar process completely absent, with the poststernite low and nearly flat in profile. The new genus differs from the males of all known Cryptopone by the following: (2) head broader than long, excluding the compound eyes (vs. head narrower than long); (3) mandibles elongate (vs. short); (4) mandibles lobate (vs. spiniform); (5) mesospiracular sclerite evidently absent (vs. this sclerite present); (6) meso- and metatibiae with no spur and one spur, respectively (vs. two spurs each); and (7) petiolar peduncle long, about as long as node is tall (vs. peduncle short, considerable shorter than height of node). Remarks. The identity of Cryptopone is significantly clarified by the phenomic data from the new species and the phylogenomic revision of Branstetter and Longino [50]. Prior to this work, the diagnostic importance of the absent subpetiolar process was obscured by the inclusion of Wadeura guianensis in Cryptopone. Now it is clear that the absence of the subpetiolar process is a unique condition among extant Ponerinae that is shared between † Desyopone gen. nov. and Cryptopone, and thus constitutes a reasonable autapomorphy within the subfamily for the two genera. No known Cryptopone, however, matches the diagnostic character combination of † Desyopone gen. nov., with conditions 3–7 being apomorphic. Critically, the elongate peduncle of † D. hereon gen. et sp. nov. is nearly unique among Ponerinae; this condition is similarly derived in Harpegnathos and is approached by Dinoponera, some Odontomachus (e.g., O. chelifer, O. coquereli), and Platythyrea (although node at middle of segment rather than posterior)., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on page 8, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409, {"references":["50. Branstetter, M. G.; Longino, J. T. UCE phylogenomics of New World Cryptopone (Hymenoptera: Formicidae) elucidates genus boundaries, species boundaries, and the vicariant history of a temperate - tropical disjunction. Insect Syst. Divers. 2022, 6, 1 - 23. [CrossRef]"]}

Subfamily Aneuretinae Emery, 1913. Type genus. Aneuretus Emery, 1893. Diagnosis (all adult castes). All adults of Aneuretinae possess the following diagnostic plesiomorphies (1–6): (1) mandibles shovel-shaped (=“triangular”) [plesiomorphy of Poneroformicia, or at least Doryloformicia]; (2) meso- and metatibiae with one spur each [synapomorphy of clade Dolichoderomorpha]; (3) petiole with complete tergosternal fusion [synapomorphy of clade Dolichoderomorpha]; (4) petiole with an elongate anterior peduncle [possible synapomorphy of clade Myrmechoderines]; (5) helcium infraaxial [possible synapomorphy of clade Myrmechoderines]; and (6) abdominal segment IV unconstricted [synapomorphy of clade Dolichoderomorpha]. Males and queens of Aneuretinae share the following diagnostic traits (7–12): (7) near-complete fore wing venation, with only the subdiscal cell open; (8) crossvein 2rs-m furcal to post-furcal, i.e., 2r-sm meeting Rs at or distad 2r-rs; (9) well-prefurcal fore wing crossvein cu-a, i.e., cu-a meeting M+Cu proximad the split of M+Cu by more than one of its own lengths; (10) absence of free M in the hind wing after the juncture of rs-m and Mf1, i.e., the abscissa between Sc+ R +Rs and M+Cu linear, without a kink; (11) hind wing anal cell short, its length less than half that of the basal cell; and (12) absence of the hind wing jugal lobe. Males of Aneuretinae have the following diagnostic plesiomorphy (13): (13) genital gonocoxa and gonostylus not strongly differentiated in size, with the dorsal gonocoxal margin continuing more-or-less evenly to that of the gonostylus. Workers and queens of Aneuretinae share the following diagnostic plesiomorphies (14, 15): (14) mandible with biseriate dentition, i.e., with small teeth interspersed between large teeth [synapomorphy of Dolichoderomorpha]; (15) basal and masticatory margins of mandible not marked, i.e., these margins curving into one another, without a distinct angle [synapomorphy of Dolichoderomorpha]. Remarks. The operational paleontological definition of Aneuretinae has relied on character states 2, 4, 6, 7 (regardless of subdiscal cell state), and 12. With the explicit recognition of character state 3—which was previously indicated for the “formicomorph subfamilies” by Bolton [11] (p. 16)—it is possible to reject the placement of † Desyopone hereon gen. et sp. nov. from the Aneuretinae. The condition of helcial axiality is here reinterpreted from Bolton [11] (p. 18), who described the helcium of Aneuretinae as “high on [the] anterior face of abdominal segment III”, which is interpretable as supraaxial sensu Keller [46]. Although the helcial tergite of worker Aneuretus is dorsoventrally short, it can be seen that the helcium is at the ventralmost position of the sternum, which does not broaden. An axial helcium is confirmed for the Baltic amber taxa † Paraneuretus and † Protaneuretus as well. Wing venation was observable for Aneuretus and † Paraneuretus. Finally, we recognize the mandibular character states 14 and 15 as critical for the identification of Aneuretinae. The states of the mandibles have not been previously remarked upon, but along with the conformation of the clypeus (not defined here), they form the gestalt of the Aneuretinae and Dolichoderinae, which was likely used by Wheeler [7] to place † Paraneuretus and † Protaneuretus, although his justifications were not made explicit. Further refinement of the aneuretine diagnosis via comparative phenomics and traditional comparative morphology is highly desirable., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on pages 6-7, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409, {"references":["11. Bolton, B. Synopsis and classification of Formicidae. Mem. Amer. Entomol. Inst. 2003, 71, 1 - 370.","46. Keller, R. A. A phylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the poneromorph subfamilies. Bull. Am. Mus. Nat. Hist. 2011, 355, 1 - 90. [CrossRef]","7. Wheeler, W. M. The ants of the Baltic Amber. Schrift. Physikal. Okonom. Gesell. Konigsberg 1915, 55, 1 - 142."]}

Tribe Platythyreini Emery, 1901. Type genus. Platythyrea Roger, 1863. Male diagnosis. In addition to the ponerine plesiomorphies, male Platythyreini are distinguished by the following: (1) mandibles worker-like, shovel-shaped (=“triangular”) [plesiomorphy]; (2) face between antennal toruli distinctly raised, such that the antennal toruli are directed relatively laterad, and are situated dorsad a depression which may receive the antenna [synapomorphy, homoplastic in Ponerini]; (3) antennal toruli usually close to or indenting the posterior margin of clypeus, toruli never distant from the posterior margin by more than half of one of their diameters [apomorphy]; (4) meso- and metatibiae with two spurs each [plesiomorphy]; (5) jugal lobe present [plesiomorphy]; (6) helcium not distinctly infraaxial, being axial to more-or-less axial, i.e., at about or just slightly below the midheight of abdominal segment III [plesiomorphy]; (7) abdominal segment IV with cinctus [plesiomorphy]; (8) cuticle pruinose [synapomorphy, homoplastic in Ponerini and some Proceratiinae]. Remarks. Because male Platythyreini have never been explicitly diagnosed, we found it necessary to provide a diagnosis in order to confirm the identification of the fossils in question. We observe that the mandibular form, tibial spur count, and cuticular sculpture of male Platythyreini are sufficient for identification at the global scale., Published as part of Boudinot, Brendon E., Richter, Adrian K., Hammel, Jörg U., Szwedo, Jacek, Bojarski, Błażej & Perrichot, Vincent, 2022, Genomic-Phenomic Reciprocal Illumination: Desyopone hereon gen. et sp. nov., an Exceptional Aneuretine-like Fossil Ant from Ethiopian Amber (Hymenoptera: Formicidae: Ponerinae), pp. 1-19 in Insects 73 (796) on pages 7-8, DOI: 10.3390/insects13090796, http://zenodo.org/record/7046409

Insects 13(9), 796 (2022). doi:10.3390/insects13090796, Fossils are critical for understanding the evolutionary diversification, turnover, and morphological disparification of extant lineages. While fossils cannot be sequenced, phenome-scale data may be generated using micro-computed tomography (µ-CT), thus revealing hidden structures and internal anatomy, when preserved. Here, we adduce the male caste of a new fossil ant species from Miocene Ethiopian amber that resembles members of the Aneuretinae, matching the operational definition of the subfamily. Through the use of synchrotron radiation for µ-CT, we critically test the aneuretine-identity hypothesis. Our results indicate that the new fossils do not belong to the Aneuretinae, but rather the Ponerini (Ponerinae). Informed by recent phylogenomic studies, we were able to place the fossils close to the extant genus Cryptopone based on logical character analysis, with the two uniquely sharing absence of the subpetiolar process among all ponerine genera. Consequently, we: (1) revise the male-based key to the global ant subfamilies; (2) revise the definitions of Aneuretinae, Ponerinae, Platythyreini, and Ponerini; (3) discuss the evolution of ant mandibles; and (4) describe the fossils as †Desyopone hereon gen. et sp. nov. Our study highlights the value of males for ant systematics and the tremendous potential of phenomic imaging technologies for the study of ant evolution., Published by MDPI, Basel

Using a fossilized birth-death model, a new phylogeny of the superfamily Evanioidea (including ensign wasps, nightshade wasps or hatchet wasps) is proposed with estimates of divergence times for its constitutive families, also corroborating the monophyly of Evanioidea. Additionally, our Bayesian analyses demonstrate the monophyly of †Anomopterellidae, †Othniodellithidae, †An-dreneliidae, Aulacidae, Gasteruptiidae, and Evaniidae, while the †Praeaulacidae and †Baissidae ap-pear to be paraphyletic lineages. Vectevaniavetula and Hyptiogastriteselectrinus are transferred in the Aulacidae. We estimate the divergence of Evanioidea in the Late Triassic (~203 Ma). Addition-ally, three new othniodellithid wasps are described and figured from mid-Cretaceous Burmese am-ber as Keradellithabasilici Jouault, Marechal, Wang & Perrichot, gen. et sp. nov., Keradellithaanubis Jouault, Marechal, Wang & Perrichot, sp. nov., and Keradellithakirina Marechal, Jouault & Perrichot sp. nov. We also document a temporal shift, in relative specific richness, between Ich-neumonoidea and Evanioidea.

Eupsenella sp. (Figs 1, 2) Material. Specimen UCM 58339 of unknown sex, part. Right lateral aspect of the body, with legs, left hind wing and parts of the antennae missing. Locality and horizon. Denson Site, Green River, Colorado, USA; Lake Uinta, Piceance Basin, Parachute Creek member; upper Ypresian, Eocene, ca. 48 Ma. Description. Body dark brown, not especially flattened, stout (length 3.23 mm). Head ovoid, apparently longer than wide; LH: ca. 0.67 mm; WH: ca. 0.36 mm; HE: 0.21 mm; compound eyes elliptical, longer than high; antennae short, only scape and two apicalmost flagellomeres preserved, apical flagellomere tapering at apex; vertex corners rounded. Mesosoma shorter than metasoma (length 1.04 mm). Fore wing micropubescent, without apparent colour patterns (LFW: 1.91 mm); C, Sc+R, M+Cu, 1Rs&1M, Rs+M, Rs and R1 preserved and tubular; 1Rs&1M angulate at junction with Rs+M; Rs apically angulate toward wing margin; poststigmal abscissa of R1 apically arched; [C], [R], [1M] (areolet), [1R1] and [2R1] cells closed; [1M] cell small, subpentagonal, aligned basally with [1R1] cell; [1R1] cell elongate, 1.0 × [2R1] cell; [2R1] cell 0.19 × LFW, evenly wide, as wide as [1R1] cell; pterostigma well-defined, elongate, slightly shorter than [1R1] cell, rectangular. Hind wing hyaline, without apparent venation. Metasoma fusiform (length 1.52 mm); segments not discernible.

TECHNOMYRMEX SVOJTKAI PERRICHOT & ENGEL SP. NOV. (FIGS 1–3; SUPPORTING INFORMATION, FIGS S1, S 2) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 565B4980-A80E-450B-A09B-603264A91F50. Type material: Holotype NHMW-N6976, a complete worker; in Early Miocene (16–23 Mya) amber from the North Shewa Zone, Amhara region, Ethiopia; housed in the Department of Mineralogy and Petrography, Naturhistorisches Museum Wien (NHMW, Austria). Paratypes MAIG 6020, two workers missing portions of legs; in Early Miocene (16–23 Mya) amber from the South Wollo Zone, Amhara region, Ethiopia; housed in the Museum of Amber Inclusions of the University of Gdansk (MAIG, Poland). Other material: MAIG 6020, one worker missing the head, portions of legs and gaster, but similar to the associated paratypes in its size and preserved structures, particularly the subcylindrical petiole, the shape and position of metathoracic and propodeal spiracles, thus considered here as conspecific; housed in the Museum of Amber Inclusions of the University of Gdansk (MAIG, Poland). Etymology: The specific epithet honours Mr Matthias Svojtka (University of Vienna) who discovered the fossil representing the holotype and generously made it available for study. Diagnosis: The species is uniquely identified within Technomyrmex by the following combination of worker characters: (1) anterior clypeal margin with a deep V-shaped median incision; (2) inner margins of incision continuously arched, with no marked angle with broadly semicircular anterolateral clypeal margins; (3) dorsum of head with a single pair of stiff setae situated above posterior level of eyes; (4) compound eyes situated around midlength of head (EPI 59), with about 40 ommatidia; (5) palp formula 6,4; (6) mandible with six conspicuous teeth apically on masticatory margin; (7) counting from the apex, first tooth longest, third and fifth teeth subequal in length, smaller than second and fourth, third to sixth teeth each separated by one denticle, sixth tooth followed by a continuous series of smaller teeth gradually decreasing in size along masticatory margin and basal angle, becoming denticles along entire basal margin; (8) the basal angle indistinct, continuously arched between margins; (9) mesosoma devoid of erect setae; (10) propodeal dorsum as long as declivitous face, their junction broadly rounded. Description: Holotype. Body integument minutely asperous, mostly glabrous except sparse stiff setae on anterior head margin and gaster. Head only slightly longer than wide. Compound eye oval, moderately large and feebly convex, its outer margin barely surpassing the outline of sides in full-face view, with 38–40 ommatidia. Ocelli absent. Frontal carinae well distant, closer to inner margin of eyes than to each other, not surpassing posterior margin of eyes and without frontal lobe, such that antennal sockets entirely exposed although directed laterad. Antenna 12-merous, scape surpassing vertexal margin by more than one-third its length; first and terminal funicular antennomeres subequal in length, nearly twice as long as broad; all other antennomeres cylindrical, only slightly longer than broad. Vertex with an erect seta (likely paired, but upper left side of head altered) longitudinally aligned with frontal carina, in profile situated at midlength between posterior margins of eye and head. Anterior clypeal margin bordered by seven stiff, short setae inclined ventrally; posterior clypeal margin semicircular, even with posterior level of toruli. Mesosoma: in profile, mesonotal dorsal outline feebly convex in its anterior third, flat and sloped in its posterior two thirds. Mesonotum and propodeum forming a distinct angle at metanotal groove. Metathoracic spiracles raised into distinct tubercules, conspicuously breaking outline of mesosoma in lateral view, distinctly anterior to metanotal groove. Dorsal surface of propodeum flat, declivitous surface convex. In profile, propodeal spiracle at junction of lateral and declivitous surfaces, slightly above midheight of sclerite. Dorsal surface of mesosoma apparently without erect setae. Legs: mesotibia and metatibia each with one spur, that of metatibia pectinate. Metasoma: petiole more than twice as long as wide, nearly as wide as high, broadest around its midlength; ventral and dorsal surfaces feebly convex; ventral surface without lobe, with two anterior longitudinal rows of three short setae. Gaster: abdominal tergite III largest, ventral surface of its anterior projection with a groove that accommodates entire petiole. Tergites III–VI with posterior margin bordered by a row of short appressed setae barely surpassing margin. Tergite VII in dorsal view small, trapezoidal. Paratypes and other specimens: As for the holotype, although incomplete or partially concealed due to preservation, but with posterior margin of head intact, not emarginate (MAIG 6020 paratypes); palpomeres clearly exposed (MAIG 6020 paratype 1), revealing a palp formula 6,4, with maxillary palp relatively long, exceeding hypostomal margin but not reaching occiput, and labial palp short, about one-third length of maxillary palp; mesonotum constricted and elongate (MAIG 6020 additional worker). Measurements and indices: Abbreviations are explained in Material and methods. Holotype: BL ~ 2.00, HL 0.50, HW 0.49, EL 0.15, FcW 0.18, SL 0.59, WL 0.77, PL 0.30, PW 0.13, PH 0.12, GL 0.80, GW 0.55, CI 98, OI 30, EPI 59, SI 129. Paratypes and additional worker: HL 0.52–0.54, HW 0.46, EL 0.11–0.15, SL 0.52, WL 0.77, PL 0.25–0.35, PW 0.20, SI 110. Note: Amber piece MAIG 6020 also contains a fourth, partially preserved worker, which displays a similar petiole and can probably be attributed to Technomyrmex. However, only the mesosoma and petiole are preserved, and it differs by a larger size (WL ~ 2.40 vs. 0.77; PL 0.77 vs. 0.25–0.35) and the presence of a dense, short pubescence covering all sclerites; it probably belongs to another species. Among the dolichoderines, the petiole reduced to a low segment without a node or scale and overhung by the first gastral segment, which is anteriorly grooved, is shared by Technomyrmex and Tapinoma Förster, 1850. The pygidium exposed in dorsal view, thus with five gastral tergites visible, as is seen on the new fossil, warrants placement in Technomyrmex (whereas the pygidium is reflexed ventrally, exposing only four gastral tergites in dorsal view in Tapinoma; see, e.g. Bolton, 2007; Fisher & Bolton, 2016). The new species is readily recognizable by its anterior clypeal margin with a deep V-shaped incision and broadly rounded lateral margins, the almost complete lack of erect setae on the dorsum of head, mesosoma and first gastral tergite (with the exception of a single pair of setae posterior to the frontal carinae), and the propodeal dorsum that is as long as, or even slightly longer than, the declivitous surface. In all other species of Technomyrmex with a deeply incised clypeal margin, the median notch is either U-shaped or semicircular, but never V-shaped. These and other features, such as the elongate mesosoma and metathoracic spiracles raised on tubercles, suggest that Technomyrmex svojtkai may belong to the bicolor group of species as defined by Bolton (2007)., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on pages 780-782, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Bolton B. 2007. Taxonomy of the dolichoderine ant genus Technomyrmex Mayr (Hymenoptera: Formicidae) based on the worker cast. Contributions of the American Entomological Institute 35: 1 - 150.","Fisher BL, Bolton B. 2016. Ants of Africa and Madagascar: a guide to the genera. Berkeley: University of California Press."]}

RAVAVY MIAFINA FISHER, 2009 Measurements and indices: (N = 4, taken from CASENT 0078664, CASENT 0080308, CASENT0115570, CASENT0474633.) BL c. 3.33–4.46, HL 2.37–2.45, HW 1.84–1.91, HWE 2.13–2.25, EL 0.34–0.43, MSL 0.08–0.09, FcW 0.49–0.55, SL 0.074– 0.98, OLL 0.24–0.28, OIL 0.44–0.56, WL 1.30–1.67, ML 0.68–0.78, MW 0.81–0.92, FWL 3.85–4.24, PL 0.15–0.19, PW –, PH 0.28–0.31, GL 1.64–2.29, GW 0.58–0.70, CI 88–92, HWI 112–114 (122), MSI 28–32, SI 39–53, OI 14–18, EPI –, OCI 44–62, MI 104–129, PI 51–67., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on page 785, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Fisher BL. 2009. Two new dolichoderine ant genera from Madagascar: Aptinoma gen. n. and Ravavy gen. n. (Hymenoptera: Formicidae). Zootaxa 2118: 37 - 52."]}

GENUS RAVAVY FISHER, 2009 Type species: Ravavy miafina Fisher, 2009: 47, figs 6, 7b., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on page 782, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Fisher BL. 2009. Two new dolichoderine ant genera from Madagascar: Aptinoma gen. n. and Ravavy gen. n. (Hymenoptera: Formicidae). Zootaxa 2118: 37 - 52."]}

GENUS TECHNOMYRMEX MAYR, 1872 Type species: Technomyrmex strenuus Mayr, 1872: 147., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on page 780, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Mayr G. 1872. Formicidae Borneensis collectae a J. Doria et O. Beccari in territorio Sarawak annis 1865 - 1867. Annali del Museo Civico di Storia Naturale 2: 133 - 155."]}

RAVAVY GOLDMANI BOUDINOT & PERRICHOT SP. NOV. (FIGS 4–5) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 693FB2F0-817E-4BD6-B2E4-5585AFC99EA2. Type material: Holotype NIGP180512, male, housed in the Nanjing Institute of Geology and Palaeontology (NIGPAS, China); paratype IGR.ET2015/001a, male, housed in the Geological Museum of the University of Rennes (IGR, France). In Early Miocene (16–23 Mya) amber from the North Shewa Zone, Amhara region, Ethiopia. Etymology: The specific epithet honours Mr Yale Goldman (Bloomfield, Connecticut) who kindly facilitated access to the type specimens of the new species. Diagnosis: Because of the limited knowledge of male ants at the global scale, this diagnosis is broken into four parts in order to establish the identity of the treated specimens: identification at the subfamilial, tribal, generic and species levels. (I) Identifiable as Dolichoderinae by the following combination: (1) clypeal condyle large, rhomboidal (see note 1 below); (2) forewing cross-vein cu-a prefurcal, i.e. joining M+Cu proximad branching point of M and Cu; (3) helcium infraaxial; (4) abdominal segment III unpetiolated, i.e. not reduced in size relative to segment IV; (5) prora absent; (6) abdominal segment IV without cinctus; (7) gonostylus dorsoventrally short relative to gonocoxite, proximally constricted and not extending proximad gonocoxite (note 2); (8) basivolsellar process present (note 3). (II) Identifiable as Bothriomyrmecini by the following combination (note 4): (9) clypeus not extending posteriorly between toruli; (10) medial hypostomal lamina absent; (11) forewing cross-vein 2rs-m absent. (III) Identifiable as Ravavy by the following features, all of which are unique within Bothriomyrmecini (note 5): (12) mandible unidentate, spatulate with an apical spiniform tooth (vs. mandible multidentate, strapshaped to spiniform) (note 6); (13) palp formula 6,3 (vs. 6,4 or ≤ 4,3) (note 7); (14) pterostigma situated immediately proximad forewing midlength (vs. situated distad); (15) marginal cell extremely long, somewhat more than twice the length of submarginal cell 1; (16) Rf appendix, i.e. distalmost free abscissa of R, long, somewhat shorter than pterostigma; (17) discal cell subrectangular, longer proximodistally than wide anteroposteriorly and size small; (18) petiolar node broadly convex in profile view (vs. anteroposteriorly narrow and squamiform); (19) penite apices not downcurved, but rather being produced distally as long, linear processes. (IV) Distinguished from Ravavy miafina by the following (note 8): (20) smaller, BL ~ 2.57–2.34, WL ~ 0.43–0.46, FWL ~ 2.17–2.52 (vs. larger, BL ~3.33–4.46, WL ~ 1.30–1.67, FWL ~ 3.85–3.99); (21) head somewhat broader, CI ~ 122 (vs. narrower, CI ~112–114, although 122 in one specimen); (22) malar space relatively long, MSI ~ 6 and space distinctly longer than proximal width of mandible in lateral view (vs. relatively short, MSI ~ 28–32 and distinctly shorter than proximal mandibular width); (23) eye relatively long, OI ~ 40–39 (vs. eye relatively short, OI ~ 14–18); (24) ocelli relatively small, OCI ~ 38 (vs. 44–62); (25) antennomere III but not IV slightly kinked (vs. both antennomeres kinked); (26) mesoscutum relatively long, MI ~ 80 (vs. short, MI ~ 104–131); (27) scutoscutellar sulcus distinctly broad (vs. short, almost line-like); (28) petiole relatively long, PI ~ 80 (vs. short, PI ~ 51–67); (29) bristle-like setae apparently not developed on gastral sternites (vs. such setae present on all gastral sternites except the first); (30) abdominal sternum IX posterior margin broadly convex (vs. broadly emarginate); (31) gonostyli relatively long, length ~ 2–3 × height (vs. short, length ~ 1 × height); (32) gonostylar apices narrowly pointed (vs. broadly rounded); (33) ventral margin of penite more-or-less linear from base to apex (vs. base produced ventrally as a dorsoventrally long, anteroposteriorly narrow, serrate lobe); and (34) apicoventral penite processes short, length from proximodorsal inflection point ~ 1 × height (vs. long, length ~ 4 × height). Notes on diagnosis: 1. A large, rhomboidal clypeal condyle is an apparent synapomorphy of the Dolichoderomorpha, i.e. Dolichoderinae + Aneuretinae. The clypeal condyle of Formicinae and other subfamilies is variable in shape, often anteroposteriorly narrow. 2. The form of the gonopod is an apparent synapomorphy of Dolichoderinae as it is not shared with Aneuretinae. 3. Most dolichoderines have a posteroventral (ventroapical) basivolsellar process, which may be short and triangular to elongate and digitate. The process is absent in Azteca Forel, 1878 and Gracilidris Wild & Cuezzo, 2006 (sister taxa), Ochetellus Shattuck, 1992 and some Dolichoderus Lund, 1831. Formicines do not have a basivolsellar process; such a process is present convergently in the Amblyoponinae, which is a synapomorphy for that subfamily (Yoshimura & Fisher, 2012; Boudinot, 2015). 4. In their recent phylogenetic revision of the Dolichoderinae, Ward et al. (2010) provided reduction or loss of the medial hypostomal lamina as a synapomorphy of the Bothriomyrmecini that occurs in both sexes. Subsequently, Boudinot et al. (2016) recognized loss of forewing cross-vein 2rs-m as another diagnostic condition. Here, we propose the ‘short’ condition of the clypeus as an additional synapomorphy of the tribe. The clypeus in both males and females extends posteriorly between the antennal toruli in the majority of dolichoderine genera with only a few exceptions (Shattuck, 1992; Boudinot, 2015), such as the neotropicus and macro clades of Leptomyrmex Mayr, 1862 (Lucky & Ward, 2010; Boudinot et al., 2016; Barden et al., 2017). 5. Males and queens of the Bornean genus Loweriella Shattuck, 1992 are unknown in contrast to the other three bothriomyrmecine genera (Bothriomyrmex Emery, 1869a, Chronoxenus Santschi, 1919, Ravavy). Because Loweriella is the sister-group of Ravavy (Ward et al., 2010), a number of the diagnostic traits of the latter genus adduced here may be synapomorphies of the two genera. 6. Yoshimura & Fisher (2011) tentatively remarked that additional, vestigial denticles may be present in R. miafina and label a possible denticle on a dissected mandible imaged with a compound microscope. Because this apparent denticle is on the lateral mandibular margin, and as the apical tooth is a consistent feature of even ‘edentate’ mandibles (Boudinot et al., 2021), we interpret the mandible of R. miafina as strictly unidentate. 7. The 6,3 palpomere count is an apparent synapomorphy of Ravavy within the Bothriomyrmecini. Loweriella has a plesiomorphic 6,4 count, while Bothriomyrmex and Chronoxenus share a derived, reduced count which is ≤ 4,3 (Bolton, 2003; Fisher, 2009; Yoshimura & Fisher, 2011). 8. The fine anatomy and morphology of R. miafina is illustrated in Fisher (2009) and especiallyYoshimura & Fisher (2011). The latter work provided a revised diagnosis of this species plus a comparative analysis of genitalic form. Comparisons were made to the images in the literature, on AntWeb (2022), and to specimens examined at the California Academy of Sciences. Description: Integument uniformly imbricate, apparently with a more-or-less even vestiture of extremely short and appressed pubescence; bristlelike setae not visible on body with exception of tarsi, gastral tergites and gonostyli, those of gastral tergites sparse. Head roughly rectangular, narrow anterad eyes and broader posteriorly; malar space relatively broad, being longer than proximal width of mandible in lateral view; posterior head margin emarginate. Compound eye bulging, length slightly less than half head-length, subreniform with subtle emargination occurring on along ventral eye margin and with> 175 ommatidia but probably not more than 200 (complete count not possible). Ocelli small; lateral ocelli relatively wideset, separated from one another by somewhat more than three lateral ocellar lengths. Frontal carinae poorly developed. Antenna 13-merous, scape about as long as compound eye and about 3 × as long as pedicel; pedicel about twice as long as wide and slightly more than half the length of antennomere III; antennomere III slightly kinked at about midlength; antennomeres III–XIII subequal in length and about 4 × as long as wide. Ocellar sensilla (setae) not visible. Anterior clypeal margin more-orless linear; clypeal setation not observable; posterior clypeal margin convex, portion of epistomal sulcus anterad and between toruli linear, weakly concave. Mandibles spiniform, tapering to their unidentate apices, thus appearing wedge-shaped in dorsal view. Labrum deeply notched apically, sides distinctly narrowing apicomediad. Palp formula 6,3; maxillary palp relatively long, exceeding hypostomal margin but not occiput; labial palp short, about one-fourth length of maxillary palp. Prementum elongate, diamond shaped. Stipes without transverse carina. Medial hypostomal lamina absent. Pronotum short, strap-like. Mesoscutum relatively long, with maximum length greater than maximum width; notauli absent; parapsides diverging anteriorly; scutoscutellar sulcus distinctly broad; lateral mesopectal sulcus sinuate. Upper metapleural region distinct, about 4 × as long as broad; lower metapleural region indistinct. Propodeum without distinct dorsal and posterior surfaces; propodeal spiracle small, circular, set slightly below metaphragmal pit. Femora puny, thin; metafemur and metatibia slightly sinuate. Tibial spur formula 1,1. Arolia small, barely visible. Petiole elliptical in lateral view, with broadly convex tergum and sternum; node not squamiform. Abdominal tergum III with broad but shallow concavity anteriorly above helcium. Abdominal segments III–VII similar in size, unmodified. Abdominal sterna VII–VIII broadly emarginate posteriorly. Abdominal sternum IX posterior margin convex, simple. Cerci not visible. Cupula not visible. Gonocoxite length uncertain; gonocoxital apex distinctly offset from gonostylar base. Gonostylus elongate, triangular; length about 2–3 × width; apex narrowly pointed. Volsellae incompletely visible; basivolsellar process present, acute; cuspis absent; lateropenite (= digitus) apex directed ventrad at nearly right angle relative to body of volsella; dorsal margin of digitus almost lobate; digital apices narrowing, almost rod-like and curved laterad. Ventral margins of penites moreor-less linear from base to apex; dorsal margins broadly convex in lateral view; apical processes directed posterad and relatively short, length from proximodorsal inflection point ~ 1 × height. Fore and hind wing venation virtually identical to R. miafina. Jugal lobe absent. Measurements and indices: Dashes indicate metric could not be taken or calculated. Holotype: BL ~ 2.34, HL 0.54, HW 0.41, HWE 0.50, EL 0.21, MSL 0.09, FcW –, SL –, OLL –, OIL –, WL 0.91, ML 0.43, MW –, FWL 2.17, PL –, PW 0.09, PH 0.12, GL 0.86, GW 0.43, CI 92, HWI 122, MSI 6, SI –, OI 39, EPI –, OCI –, MI –, PI –. Paratype: BL ~ 2.57, HL 0.54, HW 0.41, HWE 0.48, MSL 0.09, EL 0.22, FcW 0.12, SL 0.22, OLL 0.04, OIL 0.011, WL 0.98, ML 0.46, MW 0.37, FWL 2.52, PL 0.12, PW –, PH 0.15, GL 0.93, GW 0.54, CI 90, HWI –, MSI 6, SI –, OI 40, EPI –, OCI 38, MI 80, PI 80. Note: For comparison, see the metrics for R. miafina provided below., Published as part of Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej & Szwedo, Jacek, 2022, Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota, pp. 775-791 in Zoological Journal of the Linnean Society 196 on pages 782-785, DOI: 10.1093/zoolinnean/zlac053, http://zenodo.org/record/7195057, {"references":["Forel A. 1878. Etudes myrmecologiques en 1878 (premiere partie) avec l'anatomie du gesier des fourmis. Bulletin de la Societe Vaudoise des Sciences Naturelles 15: 337 - 392.","Wild AL, Cuezzo F. 2006. Rediscovery of a fossil dolichoderine ant lineage (Hymenoptera: Formicidae: Dolichoderinae) and a description of a new genus from South America. Zootaxa 1142: 57 - 68.","Shattuck SO. 1992. Generic revision of the ant subfamily Dolichoderinae (Hymenoptera: Formicidae). Sociobiology 21: 1 - 181.","Lund PW. 1831. Lettre sur les habitudes de quelques fourmis du Bresil, adressee a M. Audouin. Annales des Sciences Naturelles 23: 113 - 138.","Yoshimura M, Fisher BL. 2012. A revision of male ants of the Malagasy Amblyoponinae (Hymenoptera: Formicidae) with resurrections of the genera Stigmatomma and Xymmer. PLoS One 7: e 33325.","Boudinot BE. 2015. Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. European Journal of Taxonomy 120: 1 - 62.","Ward PS, Brady SG, Fisher BL, Schultz TR. 2010. Phylogeny and biogeography of dolichoderine ants: effects of data partitioning and relict taxa on historical inference. Systematic Biology 59: 342 - 362.","Boudinot BE, Probst RS, Brandao CRF, Feitosa RM, Ward PS. 2016. Out of the Neotropics: newly discovered relictual species sheds light on the biogeographical history of spider ants (Leptomyrmex, Dolichoderinae, Formicidae). Systematic Entomology 41: 658 - 671.","Mayr G. 1862. Myrmecologische Studien. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 12: 649 - 776.","Lucky A, Ward PS. 2010. Taxonomic revision of the ant genus Leptomyrmex Mayr (Hymenoptera: Formicidae). Zootaxa 2688: 1 - 67.","Barden P, Boudinot BE, Lucky A. 2017. Where fossils dare and males matter: combined morphological and molecular analysis untangles the evolutionary history of the spider ant genus Leptomyrmex Mayr (Hymenoptera: Dolichoderinae). Invertebrate Systematics 31: 765 - 780.","Emery C. 1869 a. Descrizione di una nuova formica italiana. Annuario del Museo Zoologico della Reale Universita de Napoli 5: 117 - 118.","Santschi F. 1919. Fourmis du genre Bothriomyrmex Emery (Systematique et moeurs). Revue Zoologique Africaine 7: 201 - 224.","Yoshimura M, Fisher BL. 2011. A revision of male ants of the Malagasy region (Hymenoptera: Formicidae): key to genera of the subfamily Dolichoderinae. Zootaxa 2794: 1 - 34.","Boudinot BE, Moosdorf OTD, Beutel RG, Richter A. 2021. Anatomy and evolution of the head of Dorylus helvolus (Formicidae: Dorylinae): patterns of sex- and caste-limited traits in the sausagefly and the driver ant. Journal of Morphology 282: 1616 - 1658.","Bolton B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.","Fisher BL. 2009. Two new dolichoderine ant genera from Madagascar: Aptinoma gen. n. and Ravavy gen. n. (Hymenoptera: Formicidae). Zootaxa 2118: 37 - 52."]}

Perrichot, Vincent, Boudinot, Brendon E., Engel, Michael S., Fls, Xu, Chunpeng, Bojarski, Błażej, Szwedo, Jacek (2022): Ants (Hymenoptera: Formicidae) from Miocene Ethiopian amber: filling gaps in the geological record of African terrestrial biota. Zoological Journal of the Linnean Society 196: 775-791, DOI: 10.1093/zoolinnean/zlac053

International audience; A new specimen of the species Litholingia rhora Ren, 2002, from the Middle-Upper Jurassic deposit of Daohugou, is used to redescribe the species and precise its diagnosis. We also propose a series of wing venation drawings highlighting the main characteristics of the wing venation all Litholingia species. Finally, we summarize the current diversity of the lacewing family Grammolingiidae.

Genus Litholingia Ren, 2002 Type species. Litholingia rhora Ren, 2002 p. 57. Included species. Litholingia rhora Ren, 2002 (type species); other species: Litholingi a eumorpha Ren, 2002, Litholingia longa Khramov, 2012, Litholingia polychotoma Ren, 2002, Litholingia ptesa Shi et al., 2011., Published as part of Jouault, Corentin, Legendre, Fr��d��ric, Perrichot, Vincent, Condamine, Fabien L. & Nel, Andr��, 2022, Redescription of Litholingia rhora Ren, 2002 (Neuroptera: Grammolingiidae) from the Middle Jurassic of Daohugou, pp. 20-26 in Palaeoentomology 5 (1) on pages 21-22, DOI: 10.11646/palaeoentomology.5.1.3, http://zenodo.org/record/6280619, {"references":["Ren, D. (2002) A new lacewing family (Neuroptera) from the Middle Jurassic of Inner Mongolia, China. Insect Science, 9, 53 - 67. https: // doi. org / 10.1111 / j. 1744 - 7917.2002. tb 00172. x","Khramov, A. V. (2012) The new fossil lacewings of Grammolingiidae (Neuroptera) from the Jurassic of Central Asia and Mongolia, with notes on biogeography of the family. Zootaxa, 3478 (1), 297 - 308. https: // doi. org / 10.11646 / zootaxa. 3478.1.28","Shi, C. F., Yang, Q. & Ren, D. (2011) Two new fossil lacewing species from the Middle Jurassic of Inner Mongolia, China (Neuroptera: Grammolingiidae). Acta Geologica Sinica, 85, 482 - 489. https: // doi. org / 10.1111 / j. 1755 - 6724.2011.00416. x"]}

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

Derolathrus capdoliensis n. sp. (Figs 2-5) urn:lsid:zoobank.org:act: 956059B5-71DF-457C-B8D8-49C8BB3F4ED6 HOLOTYPE. — IGR.CDL-2.42, slightly distorted specimen, sex indeterminate (Fig. 2). Deposited in the Geological Department and Museum of the University of Rennes, France. ETYMOLOGY. — Derived from ‘ Capdolium ’, the latin name of the type locality Cadeuil. TYPE LOCALITY. — Charentese amber; lithological horizon A1b from Cadeuil quarry at Sainte-Gemme, Charente-Maritime department, France. AGE. — Early-Late Cretaceous, latest Albian-earliest Cenomanian. DIAGNOSIS. — Head subglabrous, without deep setiferous punctures. Antennae 11-segmented, apical two antennomeres not completely fused. Antennomere 10 slightly broader and shorter than the following segment, antennomere 11 pointed apically. Eyes large, with c. 30 ommatidia, their diameter slightly greater than the distance from their anterior margin to the antennal insertions. Pronotum rather parallel-sided, broadest in the anterior quarter. Pronotal disc with an oval medial depression with a raised longitudinal groove. Femora distinctly clavate. DESCRIPTION Body minute, subcylindrical, narrowly elongate. Length from clypeus to apex of the preserved part of abdomen 0.76 mm, 0.22 mm wide across the broadest point of the elytra. Body subglabrous dorsally, dark brown to black (Figs 2; 3). Head oval, 0.2 mm long from clypeus to distalmost part of the neck, abruptly constricted behind the eyes and with a distinct neck region, slightly fractured. Head surface glabrous, lacking carinae, raised ridges or deep punctures. Mandibles small, slender, curved. Maxillary palpomeres three-segmented, with apical palpomere aciculate and the penultimate palpomere distinctly larger, approximately three times the length of the following segment and broadest in the basal third. Eyes oval, somewhat protruding, multifaceted, and consisting of approximately 30 ommatidia. Antennae 11-segmented, concealed from above and inserted anterior to the eyes, separated by slightly less than the diameter of the eyes. Antennomere 1 (scape) globular, approximately as long as wide, antennomere 2 (pedicel) 0.9 times as long and 0.8 times as wide as the preceding segment, more elongate and barrel shaped; antennomeres 3-9 moniliform, gradually shortening and widening apically, antennomeres 10-11 forming a compact club; antennomere 10 0.9 times as long and 1.1 times as broad as the following segment, widest apically; 11 pointed apically (Figs 4; 5). Temples short, less than a quarter of the length of the eye. Pronotum slightly distorted, 0.19 mm long, 1.1 times longer than wide, slightly wider than the head, approximately parallel-sided. Widest in anterior quarter. Anterior pronotal angles rounded, posterior pronotal angles obtuse. Pronotal disc with an oval medial depression with a raised longitudinal groove running from the basal quarter to the anterior quarter (Fig. 5A: pd, pg). Ventral side of pronotum strongly deformed. Procoxae globular, separated by approximately a third of their diameter across the broadest point. Mesocoxae rounded, further apart than procoxae, separated by more than half of their diameter. Metaventrite 0.24 mm long, broadening posteriorly and wider than head. Scutellum not visible. Legs short, slender with the exception of the apically swollen femur. Trochanters distinctly large, trapezoidal. Femora broadened, clavate (Fig. 5B). Tibiae slender, twice as broad in apical half as at base, lacking apical spurs. Tarsus 3-segmented, third tarsomere longer than the basal two segments combined. Claws simple, empodium not apparent. Elytra incomplete, narrower than pronotum basally,smoothly broadening towards the middle, and narrowing apically. Hind wings present, venation strongly reduced, ventral margin with approximately 30 elongate setae, dorsal margin with about 30 minute setae. Abdomen poorly preserved, apical portion apparently missing. Intercoxal process of ventrite 1 broadly rounded.

International audience; Jacobsoniidae is a species-poor family of minute polyphagan beetles distributed in tropical and subtropical regions. Both extant representatives of the family as well as fossils are rare. Here we describe a new fossil species, Derolathrus capdoliensis n. sp., from latest Albian-earliest Cenomanian Charentese amber from the Cadeuil deposit, Charente-Maritime department in south-western France. The new species is defined by several unusual characters, most notably the co-joined but not fully fused two apical antennomeres. Derolathrus capdoliensis n. sp. is approximately contemporaneous with Kachin amber (burmite), filling an important geographical gap in the Mesozoic distribution of the family. The widespread distribution of jacobsoniid beetles in the Cretaceous, encompassing the Tethyan and Austral realms, indicates a more widespread distribution of the family during this time than in the present day and suggests that the current biogeographical range of Jacobsoniidae may be a result of extinction in northern regions. The new species shows remarkable similarity to extant members of the genus and provides further evidence of prolonged morphological, and probably also ecological, stasis in Jacobsoniidae since at least the Cretaceous.

International audience; A flat wasp specimen attributed to the genus Eupsenella Westwood, 1874 is described and figured from the early Eocene compressions of the Green River Formation. This fossil is the first known outside of the Old World for this genus and documents its wide distribution during the Paleogene, while it is currently confined to Australia and New Zealand. As for several other hymenopteran genera, we assume that this regression is linked to abiotic factors, i.e., the Cenozoic climate cooling, rather than to biotic factors.

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

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

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

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

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

International audience; Examination of fossil ant specimens from various private and institutional German collections of Baltic amber has revealed a new genus and species of Myrmicinae, described here as Thanacomyrmex hoffeinsorum gen. et sp. nov. The new worker morphotype shares some similarities with the coeval extinct genus Parameranoplus Wheeler, also from Baltic amber, and the extant genera Pristomyrmex Mayr and Acanthomyrmex Emery, suggesting placement of the new fossil in the Myrmecina genus-group within the tribe Crematogastrini. The new genus highlights the tremendous diversity of ants that evolved in the Palaeogene European forests and composed the first major radiation of crown-group ants.; L’examen de fourmis fossiles dans diverses collections privées et institutionnelles allemandes d’ambre balte a révélé un nouveau genre et une nouvelle espèce de Myrmicinae, décrits ici comme Thanacomyrmex hoffeinsorum gen. et sp. nov. Ce nouveau morphotype d’ouvrière partage des similitudes avec le genre éteint Parameranoplus Wheeler, également connu de l’ambre balte, et les genres actuels Pristomyrmex Mayr et Acanthomyrmex Emery, suggérant le placement du nouveau fossile dans le groupe de genres Myrmecina, parmi la tribu Crematogastrini. Le nouveau genre souligne la grande diversité de fourmis ayant évolué dans les forêts paléogènes d’Europe et constituant la première radiation majeure du groupe couronne des fourmis.

International audience; Christophus myanmarensis gen. et sp. nov., Christophus ohmkuhnlei gen. et sp. nov., and Mintara parva gen. et sp. nov., three new species and two new genera of belytine wasps (Diapriidae: Belytinae) are described and figured from males preserved in mid-Cretaceous Kachin (Burmese) amber. These new fossils enrich the fossil record of Cretaceous diapriids, summarized here, demonstrating the diversity of the family before the K/Pg extinction and suggesting an older origin for the family.

Dhagnathos autokrator sp. nov. urn:lsid:zoobank.org:act: EC8760 A 9-9 C 00-44 A 9-9311- 2EC5 DE 24 A 4 B 1. Figs. 2AeC, 3, 8 G. Etymology. The specific epithet refers to autokrator (Greek, meaning ���self-ruler���), an individual who exercises absolute power, unrestrained by superiors; in reference to the highly powerful aspect of this ant. Holotype. IGR. BU-003, alate female (Figs. 2AeC, 3EeG). Additional specimens. HA 03, XA01 and RM 1, three alate females (Figs. 3Ae3D). Horizon and locality. Upper Cretaceous, upper Albianelower Cenomanian (ca. 99 Ma); in amber from the Hukawng Valley, Kachin State, Myanmar. Diagnosis. As for the genus, by monotypy. Description (gyne). Body length ca. 14 mm. Cuticle generally smooth, without distinct sculpturing, sparsely covered by thin, long, erect setae, the head additionally densely covered by short, adpressed setae on vertex and genae. Head only slightly longer than high and wide. Vertex and posteroventral surface rounded, anterior surface relatively flat, and genae shorter than eyes and projecting anteroventrally above mandible insertion into a cheek-like lobe. Ocelli present near top of vertex, conspicuous, ocellar diameter slightly larger than width of first antennomere; interocellar distance about half of ocellar diameter. Compound eyes bulging, reniform, 2.4 as long as wide, situated posteriorly on head (EPI 440). Antennae inserted between compound eyes around their midlength, closely flanking lateral edges of clypeus; base of antenna with basal bulb exposed, inserted within thick annular torulus. Antenna geniculate, filiform; scape short, 0.5 head length, weakly arched and broadened apically; first funicular article (pedicel) very short, 0.22 scape length, less than twice as long as wide, broadened apically; flagellomeres unusually slender, funicular article II (antennomere III) about 22 as long as wide; following antennomeres gradually decreasing in length and width. Posterior clypeal margin apparently fused, while horn is the result of de novo medial margin/ridge; anterior clypeal margin broadly rounded. Clypeal horn directed upward for its basal quarter, then bent at a right angle and directed forward for remaining length; horn gently rounded apically, without expanded lobe; dorsal surface of horn convex; ventral surface emarginate, its lateral margins prominent and prolonged basally into raised frontal carinae diverging anteriorly to reach the anterior margin of head, just above insertion of mandibles. Setation of horn consisting, on ventral surface, of a dense brush of short, peg-like denticles at apex; similar peg-like denticles widely spaced and arranged in a single row on each lateral margin, and becoming progressively denser and arranged in 2e3 longitudinal rows along lateral clypeal carinae; dorsal and ventral surfaces of horn sparsely covered by thin, long, erect setae. Labrum well exposed, large, nearly trapezoid, with anterior margin convex, posterior margin slightly emarginate medially, sides unsutured to clypeus so that anterior part of labrum is apparently movable; dorsal surface of labrum rimmed laterally by a longitudinal brush of stiff, spine-like setae, also densely coated by thin, erect setae becoming progressively longer and stiffer along lateral and posterior margins. Mandibles long (MDI 97), scytheshaped, widely spaced basally and converging apically, with tips curved and acute, nearly reaching the rounded portion of the horn as preserved; basal portion linear, short; apical portion 5 as long as basal portion, curved dorsally and posteriorly, with dorsal surface concave and rimmed on each margin by row of acute teeth and thin, erect setae directed backwards; medioventral blade between basal and apical portions forming a large, isosceles, blunt tooth perpendicular to apical portion. Palps long (visible on specimen HA 03), coated dorsally in fine, tapered setae, maxillary palp with 6 segments, as long as head capsule when combined; labial palp with 5 segments. Mesosoma. Pronotal colar pronounced, concealing propleuron in dorsal view, separated from remaining pronotal dorsum by a distinct transverse ridge; pronotal dorsum strongly concave immediately anterior to ridge, nearly flat posterior to ridge; promesonotal suture deeply impressed. Mesoscutum as long as pronotum (excluding neck) in dorsal view, about as broad as long; mesoscutal dorsal outline feebly convex, with long parapsidal furrows almost reaching anterior mesonotal margin, converging posteriorly but not touching. Mesoscutellum posteriorly expanded, in dorsal view concealing median portion of metanotum; dorsal and posterior mesoscutellar surfaces concave, their junction forming a sharp angle; dorsal mesoscutellar surface with a deep, broad, transverse groove immediately posterior to scuto-scutellar suture. Metanotum medially as high as long, with posterior surface forming distinct angle with pronotal dorsum. Propodeum 1.25 as high as long, dorsal and declivitous surfaces meet at pronounced right angle, forming conspicuous ridge; dorsal surface nearly flat, declivitous surface faintly concave; propodeal spiracle slit-like, opening posteriad, at junction of propodeal dorsum and sides; metapleural gland orifice opening laterally, protected by guard setae. Legs long and robust (mostly visible on specimen HA 03); mesocoxa distinctly shorter than pro-and metacoxae; small trochantellus present on mid- and hind legs; all femora distinctly swollen in their basal half, tibiae swollen in their apical half; ventral margin of protibia apically with large calcar gently curved, protibia possessing small subapical point, and two straight, stout setae less than half as long as calcar; mesotibia apically with two long, straight, pectinate spurs, and two short, stout setae; metatibia apically with one long, pectinate spur and one long, simple spur; tarsomeres IeIV of all legs with pairs of stout setae along entire ventral surface (8e10 pairs on tI, 4e5 pairs on tII, 3 pairs on tIII, 2 pairs on tIV), and apically with 2 pairs of stout setae each flanking a spatulate spine; additionally the ventral surface of tarsomeres IeIV covered by dense brush of thin, erect setae; pretarsal claws strong, with a distinct subapical tooth. Fore wing with Rs���f2, basal portion of Rs���f3, M ���f4, and Cu1 nebulous, all other veins tubular; pterostigma elongate, ca. 6 as long as broad; a short stub of cross-vein 1r-rs present, nebulous; Rs���f1 half as long as M ���f1, both distinctly arched; Rs���f2 and Rs���f3 nearly at right angle, Rs���f2 half as long as M ���f2; 2rs-m present, situated beyond apex of pterostigma; discal and subdiscal cells pentagonal; cu-a arising from M �� Cu and proximal to M ���f1 (Cu���f1 short); vein Cu with both Cu1 and Cu2 present. Hind wing with jugal lobe present; anterior margin with 5 median and 22 distal hamuli; vein C present; vein R present, reaching distal wing margin; Rs���f1 more than twice as long as 1rs-m; cu-a arising from M �� Cu, proximal to fork of M ���f1 and Cu (Cu���f1 short); Rs���f2, M ���f2, Cu, and A ���f2 present, not reaching wing margin. Metasoma with petiole short-pedunculate, almost 0.6 as high as long; petiolar tergite a broadly convex node, with anterior surface approximately twice as long as posterior face; subpetiolar process present, in profile forming a high, transverse, lamella pointing ventrally, with anterior face concave, posterior face vertical; not fused tergosternally, suture visible; attaching broadly to gaster. Gaster elongate. First gastral tergite with helcium pronounced, forming a post-petiolar peduncle, with anterior surface behind helcium high, oblique, and dorsal surface strongly convex, short; anteriormost part of first gastral sternite with a distinct mesal process (keel) pointing anteroventrally below helcium. Second gastral segment distinctly longer than first, with presclerite largely exposed to form a deep, broad constriction between first and second gastral segments (abdominal segments III and IV). Gastral segments unfused with deep lateral suture. Following segments poorly preserved, pygidium apparently broadly acute towards sting shaft. Measurements (holotype IGR.BU-003; in mm). HL 2.50; HoL ca.1.70; EL 1.20; ocelli diameter 0.20; MDbL 0.40, MDtL 0.55, MDaL 2.00; length/width of antennomeres: I (scape) 1.15/0.16, II (pedicel) 0.26/ 0.14, III 2.16/0.10, IV 1.50/0.07, V 1.34/0.07; WL 3.85; FWL (as preserved) 6.35 (7.90 on specimen DHA 4); PL 1.84, PH 1.00, PW 0.67., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on pages 5-7, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072

Genus Protoceratomyrmex gen. nov. urn:lsid:zoobank.org:act: D219838A-DEFE-4E2C-9599- 2FFDE59970C4. Type species: Protoceratomyrmex revelatus gen. et sp. nov. Etymology. The generic name is a combination of proto - (Greek, meaning ���first���), and the most similar genus Ceratomyrmex, in reference to the very slight clypeal horn resembling an initial stage of the dramatically developed horn of other horned hell ants. The name is masculine. Diagnosis (worker). Head configuration similar to Haidomyrmex, Haidomyrmodes, Haidoterminus, Ceratomyrmex, and Linguamyrmex; head capsule tear-drop shaped, broadened posteriorly, gradually tapering anteriorly; head distinctly broad, approximately as wide as long. Mandibles scythe-like with abbreviated dorsal development and widened mandibular ���elbow��� relative to other haidomyrmecine taxa; clypeus with well-defined lateral and posterior sulci, a slight triangulate cuticular elevation present near posterior clypeal margin, flanked by elongate trigger hairs; posterolateral clypeal margins broadly rounded; frontal triangle present as a laterally flattened rectangular elevation between antennal sockets approximately equal in height to clypeal horn; eyes reduced, ocelli absent; pronotum broadened laterally; mesosoma with distinct metanotal sclerite; propodeal spiracle circular, gaping; petiole pedunculate with broadly rounded node; ventral margin of petiole unarmed; constriction present between abdominal segments III and IV with medial v-shaped projection on first gastral tergite visible dorsally., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on page 12, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072

Genus Linguamyrmex Barden & Grimaldi, 2017. Type species: Linguamyrmex vladi Barden & Grimaldi, in Barden et al., 2017: 839., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on page 13, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072, {"references":["Barden, P., Herhold, H. W., Grimaldi, D. A., 2017. A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure. Systematic Entomology 42, 837 e 846. https: // doi. org / 10.1111 / syen. 12253."]}

Genus Aquilomyrmex gen. nov. urn:lsid:zoobank.org:act: 5 AFEF 07 E- 2228-4A9 D-AEC 1- 8D2680267734. Type species: Aquilomyrmex huangi gen. et sp. nov. Etymology. The generic name is a combination of aquilex (Latin, meaning ���dowser���), and myrmex (Greek, meaning ���ant���), and refers to the dowsing stick-like clypeal and labral processes. The name is masculine. Diagnosis (gyne). Head dorsoventrally flattened, with large compound eyes situated dorsolaterally at anterior margin of head; with prominent, anteriorly protruding frontal shelf, clypeal horn and labrum. Antennae inserted laterally on frontal shelf, well in front of compound eyes and above base of clypeal horn; antennal scape long. Apex of clypeal horn bifurcated widely, with each bifurcation subsequently terminating in a bilobed pad; lateral margins of horn connecting obliquely with anteroventral angles of head, just above mandibular insertion, connected by a cleared vertical cuticle, rimmed by short thin erect setae and occasional serrations. Labrum situated ventrally of horn, with size and shape similar to horn except for apex, which is only bifid; ventral margin coated by peglike denticles on its basal half. Mandibles sickle-shaped, inserted ventrally very close to compound eyes, with all margins smooth (triangular blade absent) and acute tips converging between labral apex. Legs very long, with femora distinctly swollen basally, and apically with two flange-like cuticular lobes flanking the base of tibia. Petiole node-shaped, with small subpetiolar process. A deep, girdling constriction between first and second gastral segments., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on page 10, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072

Chonidris insolita sp. nov. urn:lsid:zoobank.org:act:3 A 8ED551-78 C 9e4 B 0F-8 E 10- 6 C 94641 F 5 A 08. Figs. 2DeF, 4, 8 F. Holotype. NIGP 171998, alate female (Figs. 2D, 4 H). Paratype. NIGP 172003, alate female (Figs. 2 E, 4FeG). Additional specimens. HA 04 and RM 2, two alate females (Figs. 2 F, 4AeE). Horizon and locality. Upper Cretaceous, upper Albianelower Cenomanian (ca. 99 Ma); in amber from the Hukawng Valley, Kachin State, Myanmar. Etymology. The specific epithet derives from insolitus (Latin, meaning ���strange���), and refers to the unusual accommodation of the mandibles within the clypeus. Diagnosis. As for the genus, by monotypy. Description (gyne). Body length around 10 mm. Cuticle smooth, densely covered by short adpressed setae; additionally with long, thin, erect setae sparsely covering head capsule, mesosoma, legs, and apical portions of gastral segments IeIII, setae more densely covering pygidium. Head about as long as broad, longer than high, circular in frontal view. Vertex and posteroventral surface strongly rounded, anterior surface flat, with short genae (ca. 0.2 eye length) projecting into cheek-like lobe anteroventrally above mandible insertion. Ocelli distinct, positioned on small, raised, triangular prominence; ocellar diameter as broad as base of first antennomere, interocellar distance about one ocellar diameter. Compound eyes bulging, oval, 0.6 as broad as long, removed from lateral margins of head capsule. Antennae inserted between compound eyes below their midlength, closely flanking lateral edges of clypeus at base of horn's stalk; base of antenna with basal bulb exposed, inserted within thick annular torulus opening obliquely dorsad; antenna elongate; scape ca. 0.8 head length, weakly arched, cylindrical; pedicel very short, 0.2 scape length, about as long as wide, slightly broadened apically; first flagellomere longest antennomere, about 8 as long as wide (assessed from paratype, where antennae are flattened and elongate but respective ratios are kept); following antennomeres gradually decreasing in length. Posterior and lateral margins of clypeus (epistomal sutures) visible, indicating the horn is the result of an elevated anterior margin. Clypeal horn short (HoI 28e30), expanded at right angle from frons, strongly arched with dorsal surface convex and ventral surface furrowed; lateral margins prolonged basally on frons into raised carinae diverging toward anterior angles of head, just above insertion of mandibles; ventral surface and lateral carinae rimmed by row of short, peg-like denticles, and long, fine, erect setae. Labrum exposed between clypeal carinae, with anterior margin broadly convex and slightly notched medially, posterior margin obscured; dorsal surface coated by long, thin, erect setae. Dorsal margin of mandibles curved dorsally immediately from base, in profile view without distinct elbow, their tips reaching nearby horn's apex, dorsal mandibular margin armed apically as series of sharp, curved teeth; largest apical tooth preceded by two medial teeth of equal length following slight gap; basal portion of mandible (from base to base of medioventral blade) short, simple; distal portion projected medially and ventrally in a large, triangular blade tapering to the tip; the blade with dorsal surface concave, ventral corner rounded, and apical half serrate; tip of mandible with a preapical tooth immediately following the blade and projected posteriorly, and one apical acute tooth curved posteriorly; when mandibles closed, their medial margins parallel and closely approximated, slightly overlapping at ventral corner of blade; accommodated in the triangular clypeal cavity (i.e., concealing the anterior clypeal margin, labrum and ventral surface of horn); maxillary palp with 5 visible segments, labial palp with 3 visible segments. Mesosoma about twice as long as high. Pronotal colar short, separated from remaining pronotal dorsum by a distinct transverse ridge; pronotum nearly vertical immediately anterior to ridge, feebly convex posterior to ridge (dorsum); posterior pronotal margin with distinct thickening dorsally. Promesonotal suture complete, appears flexible. Mesoscutum shorter than pronotum, mesoscutal dorsal outline faintly convex, with long parapsidal furrows converging posteriorly to reach anterior mesonotal margin. Mesoscutellum prominent; dorsal mesoscutellar surface convex, posterior surface slightly concave, their junction forming a sharp angle. Metanotum medially as high as long, with posterior surface forming distinct angle with propodeal dorsum. Propodeum 1.5 as high as long, dorsal and declivitous surfaces flat, separated by thin ridge, forming sharp angle; propodeal spiracle slit-like atop rounded nodule, situated around junction of propodeal dorsum and sides. Metapleural gland orifice a small circular concavity. Mesopleuron and metapleuron coated in thin, tapered setae. Legs robust, mesocoxa distinctly shorter than pro-and metacoxae, small trochantellus present on mid- and hind legs; all femora moderately swollen in their basal half; ventral margin of protibia apically with large calcar and two straight, stout setae less than half as long as calcar; mesotibia apically with two long, straight, spurs, one pectinate and one simple; additionally with three short stout setae; metatibia apically with one long, pectinate spur, one long, simple spur, and one short, stout seta; tarsomeres IeIV of all legs with pairs of short, stout setae along entire ventral surface (6 pairs on tI, 3 pairs on tII, 3 pairs on tIII, 2 pairs on tIV), and apically with 2 pairs of simple, stout setae each flanking a spatulate seta (�� plantar lobe); pretarsal claws strong, with a distinct subapical tooth and pulvilus. Wing venation as in Dhagnathos except, in fore wing, the pterostigma 3 as long as broad, cross-vein Rs���f1 not arched, short stub of 1r-rs tubular, M ���f2 extremely short (1m-cu almost arising at level of Rs���f2). Hind wing with 11 distal hamuli present, jugal lobe not visible, but lobe may be lost due to damage. Metasoma. Petiole short-pedunculate, massive, 0.84 as high as long; in profile, with posterior margin oblique and broadly attached to first gastral tergite (AIII); tergite and sternite of petiole and AIII unfused (suture distinct); petiolar tergite a strongly convex node, with anterior surface approximately twice as long as posterior surface which is nearly vertical in its anterior half, oblique in its posterior half; petiolar sternite with subpetiolar process and lamella (in profile) with a high, transverse tooth pointing ventrally, with anterior face flat, posterior face concave; process followed by a smaller but distinct triangular tooth. First gastral segment bellshaped; tergite with helcium faintly pronounced, with anterior surface behind helcium high, oblique, and dorsal surface strongly convex, short; anteriormost part of first gastral sternite with a distinct mesal process (keel) projecting anteroventrally below helcium. Second gastral segment (AIV) distinctly longer than first, with presclerite largely exposed to form a deep, broad constriction between AIII and AIV. Following segments more or less telescoped, pygidium convex, sting well exserted, distinctly arched dorsally. Measurements (in mm) (holotype NIGP 171998), [paratype NIGP 172003], {specimen RM 2}. HL [1.42] {1.68}; HoL [0.86] {0.658}; EL [0.72]; MDL [1.44] {1.10}; length of antennomeres: I (scape) [1.00] {0.96}, II (pedicel) [0.17] {0.21}, III [1.42] {0.92}; WL (3.66) {3.30}; FWL (5.2) {5.50}; PL (1.33), PH (excluding process) (1.12)., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on pages 8-10, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072

Linguamyrmex brevicornis sp. nov. urn:lsid:zoobank.org:act: D91488C6-3BEB-4A21-9F0E- 3101BB9CCA85. Figs. 6CeE, 7. Etymology. The specific epithet is a combination of brevi (Latin, meaning ‘short’) and cornus (Latin, meaning ‘horn’), and refers to the smaller clypeal horn compared to the type species L. vladi. Holotype. NIGP172001, worker (Figs. 6C, 7D). Additional specimens. LA01, alate female (Figs. 6D, 7AeB), and RM 4, worker (Figs. 6E, 7C). Horizon and locality. Upper Cretaceous, upper Albianelower Cenomanian (ca. 99 Ma); in amber from the Hukawng Valley, Kachin State, Myanmar. Diagnosis (females). Closely similar to L. vladi, but differs from this species by having a shorter clypeal horn (horn index HoI ¼ 31e 34 in L. brevicornis, 56 in L. vladi), with stalk much reduced, slightly broader than long, with the setose pad more hexagonal and aligned with stalk (circular and with a distinct bend between stalk and clypeal pad in L. vladi), and with clypeal pad coated in stout setae. It is also differentiated by the mandibles with two teeth on ventral corner of medioventral blade (teeth absent in L. vladi) and, in workers, with the apical mandibular portion shorter than basal portion (apical portion longer in L. vladi). In addition to the difference in the proportion of the mandible, the female castes of L. brevicornis differ by their size (body length ca. 7 mm in gyne, 4 mm in worker), by the antennae with flagellomeres compact in worker, more elongate in gyne, and by subpetiolar process present in worker. Description (gyne). Total length of body around 7 mm. Cuticle minutely shagreened, densely covered by adpressed, minute setae, the apical gastral segments additionally with sparse pilosity of long, fine, erect setae becoming gradually denser toward apex. Head. As in Linguamyrmex vladi, but with compound eyes slightly reniform, emarginate in middle of posterior margin. Ocelli distinct, positioned on a prominent cuticular triangle, in dorsal view, with posterior margin of lateral ocelli aligned with posterior margin of eyes. Antenna filiform, with flagellomeres elongate, cylindrical except apical one, which has acute tip; third antennomere twice as long as following one. Antennal sockets immediately flanking a medial frontal triangle that is sharply expanded anteriorly above clypeal horn, and connecting ventrally with dorsal surface of horn by a cleared, laterally flattened cuticle; frontal triangle projection terminates anteriorly at approximately same length as clypeal horn with slight indentation just dorsad clypeal horn, giving the appearance of a second, small, broadly rounded horn. Clypeal horn originating at base of frontal triangle, short and straight, in profile view almost perpendicular to longest axis of head. Horn paddle-shaped, with short, compact stalk and hexagonal, setose pad; stalk apparently glabrous, 0.10 mm in length, 0.12 mm in width; setose pad 0.26 mm in length, 0.20 mm at greatest diameter, with anterior margin straight, transverse; dorsal surface of setose pad glabrous, ventral surface densely and uniformly coated with very short, velcro-like vestiture; trigger hairs not visible. Lateral clypeal margins extending from above mandibular insertions toward horn as gently curved lines. Anterior clypeal margin broadly concave. Labrum visible. Mandible scythe-shaped; basal portion linear, short, meeting apical curved portion nearly at right angle; medioventral triangular blade bearing one large apical tooth and one small preapical tooth, with concave dorsal surface coated with pointed setae; apical portion of mandible posterior to medioventral blade 1.6 as long as blade, with tip broadly rounded. Mandibles preserved in closed position (with tips of apical portion reaching nearby clypeal setose pad), almost entirely parallel and closely approximated, except ventral corners of blades overlapping and tips slightly diverging. Maxillary palps 5-segmented, labial palps 3- segmented. Mesosoma. Pronotal colar pronounced but not concealing entire propleuron in dorsal view; pronotal dorsum weakly convex. Promesonotal suture present, complete. In dorsal view, mesoscutum as long as pronotum (excluding neck), about as broad as long, with lateral and posterior margins carinate; posterior mesoscutal margin broadly concave; dorsal mesoscutal surface with long parapsidal furrows almost reaching anterior mesonotal margin, converging posteriorly but not touching. In profile view, pronotal dorsal outline feebly convex, mesoscutal dorsal outline nearly flat. Mesoscutellum prominent posteriorly; posterior mesoscutellar surfaces concave, their junction forming a sharp angle; dorsal mesoscutellar surface with a deep, broad, transverse groove immediately posterior to scuto-scutellar suture. Metanotum medially as high as long, with posterior surface forming distinct angle with pronotal dorsum. Propodeum 1.3 as high as long; in profile view, propodeal dorsum distinctly lower than pronotal dorsum; propodeal dorsum nearly flat, declivitous surface feebly convex, their junction broadly rounded. Propodeal spiracle slit-like, oriented posteriad. Metapleural gland opening circular, gaping. Legs long and robust, mesocoxa distinctly shorter than pro- and metacoxae; small trochantellus present on mid- and hind legs; all femora distinctly swollen in their basal half, tibiae swollen in their apical half; ventral margin of protibia apically with large calcar and two simple, stout setae less than half as long as calcar; mesotibia apically with two long, straight, pectinate spurs, and two short, straight, stout setae; metatibia apically with one long, pectinate spur and one long, simple spur. Tarsomeres IeIV of all legs with pairs of stout setae along entire ventral surface (8e10 pairs on tarsomere I, 4e5 pairs on tarsomere II, 3 pairs on tarsomere III, 2 pairs on tarsomere IV), and apically with 2 pairs of stout setae each flanking a spatulate spine; additionally the ventral surface of tarsomeres IeIV covered by dense brush of fine, erect setae; pretarsal claws thick, with a distinct subapical tooth. Fore wing with veins M∙f4 and Cu1 nebulous, not reaching wing margin; remaining veins tubular; cross-vein 1r-rs absent; Rs∙f1 faintly arched; M∙f1 distinctly arched, twice as long as Rs∙f1; Rs∙f2 nearly at right angle with Rs þ M, half as long as M∙f2; Rs þ M and M∙f2 not aligned, so that discal cell pentagonal, with vein 1m-cu distinctly longer than M∙f1; second submarginal cell long, crossvein 2rs-m situated far beyond apex of pterostigma; cross-vein cu-a arising from Cu, shortly distal to M∙f1. Hind wing with jugal lobe present. Metasoma. Petiole short-pedunculate; petiolar tergite forming a broadly convex node, with anterior surface approximately twice as long as posterior surface; subpetiolar process present, in profile a high, transverse, tooth pointing ventrally, with anterior surface concave, posterior surface vertical; transverse sulcus visible across petiole near basal margin. First gastral segment with helcium pronounced, forming a post-petiolar peduncle, with anterior surface posterior to helcium high, oblique, and dorsal surface strongly convex, short; anteriormost part of first gastral sternite with a distinct mesal process pointing anteroventrally below helcium. Second gastral segment distinctly longer than first, with presclerite largely exposed to form a deep, broad constriction between first and second gastral segments (abdominal segments III and IV). Posterior margin of gastral segments II onward with sparse, elongate setae. Pygidium nearly glabrous while hypopygium with sparse setae; pygidium apparently broadly acute towards sting shaft. Wingless female (worker). Smaller than gyne (total length of body around 4 mm), and with the following differences: compound eyes oval, ocelli faintly visible, not positioned on a prominent cuticular triangle. Antenna more compact, with flagellomeres IIeIX gradually broadened apically, apical flagellomere with rounded apex. Clypeal setose pad more quadrangular, the ventral surface with a pair of trigger hairs originating at pad base. Mandible with medioventral triangular blade larger and apical portion (posterior to blade) reduced, 0.6 as long as blade. Mesosoma and metasoma as in L. vladi except propodeum higher than long, petiole with subpetiolar process present, and girdling constriction deeper between abdominal segments III and IV (AIII with an oblique posterior surface). Protibia with one large apical spur (calcar) and two short stout setae; mesotibia with two simple spurs and three conspicuous setae positioned along anterior margin of tibial apex; metatibia with one pectinate and one simple spur, additionally with a brush of 8e10 setae flanking the pectinate spur. Sparse, erect setae positioned on dorsum of pronotum, propodeum, petiolar tergite, and first and second gastral tergites and sternites. Pygidium not preserved. Measurements (in mm) (worker holotype NIGP 172001) [worker specimen RM 4] {gyne specimen LA01}. HL [0.82] {0.82}; EL [0.21] {0.42}; HoL [0.17] {0.36}; MDL [0.59] {0.80}; length of antennomeres: I (scape) (0.57) [0.67] {1.04}, II (pedicel) (0.09) [0.12] {0.14}, III-XII (0.35, 0.17, 0.16, 0.15, 0.15, 0.13, 0.13, 0.12, 0.13, 0.17) {0.80, 0.40, 0.34, 0.32, 0.31, 0.30, 0.30, 0.32, 0.32, 0.35}; WL [1.29] {2.29}; PL {0.70}.

Genus Chonidris gen. nov. urn:lsid:zoobank.org:act:664 B 304Ce41 C 9e4 F 39-9561- 2 B 055 E 815 C 3 B. Type species: Chonidris insolita gen. et sp. nov. Etymology. The generic name is a combination of chon�� (Greek, meaning ���funnel���) and idris (Greek, meaning ���ant���), in reference to the shape of the clypeus. The name is feminine. Diagnosis (gyne). Highly similar to Dhagnathos, but smaller (body length ca. 9 mm), with clypeus triangulate and expanded posteriorly into rounded horn, lateral margins of clypeus raised into serrated carina; propodeum with posterodorsal ridge; ventral margin of petiole with pointed spicule-like process and lamella; gastral segments I and II (abdominal segments III and IV) with conspicuous constriction. Separable from Dhagnathos with clypeal horn, apical portion of mandibles, and flagellomeres distinctly more compact; with inner margin of mandible projecting medioventrally into a large triangular blade that is gradually tapering to the tip of mandible; this blade with ventral corner rounded and dorsomedial margin serrate; in frontal view, when mandibles closed, ventral corners of blades slightly overlapping and medial margins almost parallel, nearly touching, so that labrum and clypeal area ventral to horn are entirely concealed, and mandibles are encased in clypeal triangle., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on page 7, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072

Protoceratomyrmex revelatus sp. nov. urn:lsid:zoobank.org:act: 02E35429-75CB-45F0-8446- AE4598E48E21. Figs. 6AeB, 8B. Holotype. NIGP172002, worker (Figs. 6AeB). Horizon and locality. Upper Cretaceous, upper Albianelower Cenomanian (ca. 99 Ma); in amber from the Hukawng Valley, Kachin State, Myanmar. Etymology. The specific epithet derives from revelatus (Latin, meaning ‘reveal’ or ‘show’), and refers to the clypeal margins, horn, and frontal triangle, which informed interpretations of morphological development in hell ants. Diagnosis. As for the genus by monotypy. Description (worker). Total length 4.30 mm. Cuticle generally glabrous throughout. Head. Vertex broad, gradually rounded posterolaterally, medially flattened; head capsule tapered gradually toward anterior margin with maximum width at vertex approximately 2 that at mandibular insertion; head flattened anteriorly, with steep elevational incline present dorsally to oral opening; ventral surface of head severely depressed around occipital foramen. Ocelli absent, oval-shaped eyes reduced, positioned near midlength of head in lateral view, with dorsal margin abutting vertex of head. Mandibles scythe-like, laterally flattened, dorsoventrally expanded, appearing broad in lateral view; medial margin of mandibles slightly bowed anteriorly, producing cup-like curvature; dorsally developed apical tooth arising gradually, producing curved dorsal mandibular margin from basal arm of mandible, triangulate blade present anteroventrally at “elbow” junction of basal mandibular margin and apical tooth, fine setae present on the lateral margin of this expansion; dorsal margin of mandible unarmed; length of apical tooth and basal margin of mandibles approximately equal. Maxillary palps elongate, ca 0.75 length of head capsule, comprising six equally sized palpomeres; labial palps stout, roughly equal in length to two maxillary palps, comprising four equally sized palpomeres. Clypeus elongate and steeply elevated, sclerite is well defined by sulci, posterior and lateral margins meet broadly as gradually rounded suture; small, triangulate clypeal hornpresent, slight anterior widening visible from oblique view; long, fine seta present at base of clypeal horn, approximately 2 length of horn itself, reaching beyond apex of mandibles as preserved (second, symmetrical hair flanking horn presumably lost). Antennal sockets present just dorsad posterior margin of clypeus; antenna with scape elongate, third antennomere more than twice as long as following one. Cuticle raised between antennal sockets into laterally flattened projection, likely homologous with “frontal triangle” of other haidomyrmecine taxa; projection with sharp anterior face, slightly declined dorsal face, and gradually sloping posterior face, 0.23 mm in length and 0.08 at greatest height. Mesosoma. Pronotum and propleuron anteriorly extended into neck to meet and accommodate depression of head; propleuron reduced in lateral view, only faintly visible; pronotum broadly arched and dome-like; expanded ventrally in lateral view, with maximum height approximately equal to pronotal length in dorsal view; posterior margin of pronotum demarcated by well-defined mesonotum, resulting in a sharply circular posterior margin; in lateral view, posterior margin of pronotum nearly extending to posterior margin of mesonotum. Maximum height of pronotum, mesonotum, metanotum, and propodeum all approximately equal, with metanotum and propodeum slightly lower in elevation. Propodeum height and length approximately equal, broadly rounded dorsally; large propodeal spiracle situated high, circular and gaping; metapleural gland opening slightly oval-shaped. Procoxa approximately twice as long as mid- and hind coxae; femur and tibia of each leg coated in fine setae; protibia with large calcar and two stiff setae approximately 0.33 length of calcar; mid- and hind legs with two tibial spurs of equal size; trochantellus present on mid- and hind legs; tarsomeres with fine, stiff setae on underside; conspicuous pretarsal claw present. Metasoma. Petiole longer than high, node-shaped, coated in short, fine setae; peduncle short, comprising approximately 0.2 length of petiole itself; petiole node broadly rounded, with anterior surface gradually increasing in elevation, slightly flattened dorsally; posterior surface of petiole attaches to gastral segment I (abdominal segment III) broadly, following a slight decrease in elevation; ventral margin of petiole appears unarmed, without any process or tooth. Abdominal segment III with significant helcium, approximately equal in length to petiole peduncle; sternite possessing a slight ventral keel, triangulate and projecting just ventral to petiole as preserved. Darkened lateral sulci visible on each gastral segment, present along lower one-third of the gaster. Deep constriction present as circular banding between abdominal segments III and IV, this circular constriction interrupted by slight vshaped posterior expansion of abdominal tergite III, visible from above. Pygidium with elongate, tapered setae; third valvula visible above sting; sting curved near apex. Measurements (in mm) (holotype NIGP 172002). HL 0.82; EL 0.20; HoL ca. 0.09; MDbL 0.36, MDtL 0.09, MDaL 0.41; length of antennomeres: I (scape) 0.42, II (pedicel) 0.08, IIIeXII 0.27, 0.13, 0.14, 0.12, 0.08, 0.10, 0.10, 0.11, 0.10, 0.12; WL 1.21; PL 0.33, PH 0.25.

Aquilomyrmex huangi sp. nov. urn:lsid:zoobank.org:act: 9C026315-9D66-406B-BF2E- 401F12305A2D. Figs. 2GeH, 5, 8 C. Holotype. NIGP 172000, alate female (Figs. 2 G, 5GeH). Paratype. NIGP 171999, alate female (Figs. 5eF). Additional specimens. HA 06 and RM 3, two alate females (Figs. 5AeD). Horizon and locality. Upper Cretaceous, Lower Cenomanian (ca. 99 Ma); in amber from the Hukawng Valley, Kachin State, Myanmar. Etymology. The specific epithet is a patronym honoring Mr. Huang Yiren who provided four specimens for this study. Diagnosis. As for the genus, by monotypy. Description (gyne). Estimated body length around 9 mm. Cuticle smooth, densely covered by minute, adpressed setae, the pygidium additionally with dense, fine, erect setae. Head prognathous, dorsoventrally flattened; in dorsal view, with lateral margins slightly converging posteriorly, occipital corners broadly rounded, posterior margin straight. Vertex elongate, nearly flat. Ocelli forming a small, weakly prominent triangle situated on vertex far posteriorly to posterior eye margins. Compound eyes large, oval, with outer margin only feebly extending beyond side of head laterally, situated dorsolaterally near anterior margin of head. Genae much reduced. Antennae inserted anteriorly to anterior margin of compound eyes, dorsally to base of clypeal horn, and immediately flanking lateral edges of a prominent frontal shelf; frontal shelf is distributed into a three-pronged projection comprising a median membranous support stalk, flanked by strong, turreted toruli partially concealing antennal bases in dorsal view; membranous stalk is flattened laterally, appearing as a slender line in dorsal view; in lateral view, stalk appears broad with conspicuous medial hole (potentially arising from desiccation; however, present in holotype and paratype), lower portion of stalk with ventral margin fused to the dorsal margin of the clypeal horn. Base of antenna with basal bulb exposed. Antenna geniculate, 12- segmented; scape long (SI 140), weakly arched in its basal half; pedicel short, 0.10 scape length, less than twice as long as wide, broadened apically; antennomere III and XII longest flagellomeres, AIII cylindrical, about 3.6 as long as wide, following flagellomeres gradually (weakly) increasing in width, apical flagellomere with rounded apex. Clypeal horn dorsoventrally flattened, protruding anteriorly (HoI 50), gently curved dorsally toward apex; with dorsal surface convex in profile view, ventral surface apparently concave; terminus of membranous horn bifurcated widely, with each bifurcation terminating in a subsequent bilobed pad; each pad with a small, medial, rounded lobe and a larger, lateral, rounded lobe, with tapered setae present along these lobes; lateral margins of horn connecting obliquely with anteroventral angles of head, just above mandibular insertion, by a cleared vertical cuticle, rimmed by short, thin, erect setae and occasional serrations. Labrum situated ventrally to horn, with to horn in size and shape except for apex only bifid (not each bifurcation bilobed; i.e., with the form of a snake tongue); ventral surface coated by short, darkened, thick, peg-like denticles on its basal half, additionally with sparse, long, fine, erect setae throughout. Mandibles sickle-shaped, inserted very close to compound eyes, lateral to hypostoma in ventral view; weakly spaced basally and converging apically, with tips acute and reaching to horn's apex between anterior median notch of labrum and clypeal horn; basal portion linear, directed ventrally; apical portion about 3 as long as basal portion, curved dorsally approximately at 45 from basal portion, with dorsal surface apparently concave and smooth. Palps short, not reaching to occipital margin; maxillary palp with 5 visible segments, labial palp with 3 visible segments. Mesosoma elongate, distinctly longer than high and wide. Pronotal neck pronounced, about as long as wide, 0.4 as wide as maximal width of pronotal dorsum; pronotal dorsum distinctly lower than remaining mesosomal dorsum, with posterior margin strongly concave. Promesonotal suture present, complete. Mesoscutum in dorsal view shorter than pronotum (excluding neck), shorter than broad, with a distinct oblique anterior face forming a sharp angle with pronotal dorsum. Mesoscutal dorsal outline strongly convex, with faint parapsidal furrows converging posteriorly, almost reaching posterior mesoscutal margin. Mesoscutellum anteriorly flat, posteriorly strongly convex; anterior half bordered on each side by sharp carina converging posteriorly toward posterior convexity. Metanotum medially exposed in dorsal view, about half as long as mesoscutum. Propodeum convex, about as high as long, dorsal and declivitous surfaces forming a continuous curve; propodeal spiracle slit-like, situated high on sides, opening posteriad. Metapleural gland orifice a circular concavity at posteroventral margin of propodeum. Legs very long, slender. Mesocoxa only slightly shorter than procoxa, half as long as metacoxa. Small trochantellus present on mid- and hind legs. All femora distinctly swollen in their basal half, more than twice as broad as apical portion; ventral margin of each femur apically with two flange-like cuticular lobes flanking the base of tibia, these lobes increasingly larger from fore-to hind legs. Tibiae gradually increasing in width apicad. Dorsal surface of mid- and hind tibiae with a longitudinal row of 18e20 short, erect, stiff setae, and a paired row of long, erect, fine setae. Ventral margin of protibia apically with large calcar gently curved and one stout seta less than half as long as calcar; additionally the dorsal margin with three small, stiff setae. Mesotibia apically with two small, simple spurs; dorsal margin with 4 short, stiff setae. Metatibia apically with one long, pectinate spur and one short, simple spur; dorsal margin with 2 stiff setae. Tarsomere I of all legs elongate, longer than combined length of following tarsomeres IIeV; tarsomere I of fore leg coated with longitudinal row of stiff setae on dorsal margin, and dense, short, fine, erect setae throughout; tarsomere I of hind leg very long, 0.8 as long as tibia. Pretarsal claws with a distinct subapical tooth. Fore wing with veins M∙f4 and Cu1 nebulous, not reaching wing margin; remaining veins tubular; cross-vein 1r-rs absent; Rs∙f1 faintly arched; M∙f1 distinctly arched, twice as long as Rs∙f1; Rs∙f2 nearly at right angle with Rs þ M, half as long as M∙f2; Rs þ M and M∙f2 not aligned, so that discal cell pentagonal, with vein 1m-cu distinctly longer than M∙f1; second submarginal cell long, crossvein 2rs-m situated far beyond apex of pterostigma; cross-vein cu-a arising from Cu, shortly distal to M∙f1. Hind wing with 14 distal hamuli, jugal lobe not visible, base of hind wing lost. Metasoma. Petiole short-pedunculate, twice as long as broad; petiolar tergite a broadly convex node, with anterior and posterior surfaces subequal in length; posterior surface oblique in its basal half, horizontal in its distal half; broadly attached to base of first gastral tergite; subpetiolar process reduced to a small, laterally flattened rectangle present ventrally to petiole node ascent. First gastral segment bell-shaped; first tergite with helcium pronounced, forming a post-petiolar peduncle, with anterior surface posterior to helcium high, oblique, and dorsal surface strongly convex, short. Deep, girdling constriction between first and second gastral segments (abdominal segments III and IV). Second gastral segment distinctly longer than first, with deep lateral sulcus. Dorsum and lateral margins of pygidium coated in dense, tapered setae, hypopygium with long, sparse setae at terminus. Third valvula present. Sting present, strong. Measurements (in mm) (holotype NIGP 172000), [specimen RM 3]. HL (1.40) [1.33], HW (1.05), Hh (0.50); HoL (0.86); EL (0.56 in dorsal view); MDL (ca. 1.00); length of antennomeres: I (scape) (1.47) [1.80], II (pedicel) (0.16) [0.18], III-XII (0.25, 0.18, 0.16, 0.13, 0.13, 0.16, 0.13, 0.13, 0.13, 0.25); WL [3.50]; PL (0.61).

Genus Dhagnathos gen. nov. urn:lsid:zoobank.org:act: 48DA68FC-1DF6-4 A 90-86DA- 4EF527D762 F 4. Type species: Dhagnathos autokrator sp. nov. Etymology. The generic name is a combination of Dha, a singleedged sword with long, gently curved blade common throughout mainland Asia, and often called ���the national sword of Burma���, and gnathos (Greek, meaning ���jaw���), in reference to the mandibles' shape. The name is masculine. Diagnosis (gyne). Large, robust ant, body length ca. 14 mm. Clypeus funnel-shaped resulting from extreme posterior expansion, extends well beyond antennal insertions, with lateral margins raised into carinae arising above mandible insertion and converging posteriorly to form a clypeal horn; horn bent forward at right angle from frons, its tip gently rounded, not spatulate, its underside deeply furrowed, forming a channel opening toward the labrum; clypeal carinae, including horn's edges, rimmed by stout, short, tooth-like denticles. Mandibles scythe-like, the elbow between basal and distal portions with a strong (isosceles) triangular blade pointing medially and ventrally, the apical portion long, curved upwards and backwards, acutely tapering to the tip and with dorsal (posterior) margin furrowed and serrated; mandibles widely spaced, approximated only apically, with medioventral blade not overlapping (in frontal view, when closed, with distal portions aligned with frontal carinae so that the labrum and clypeal area below horn are exposed). Labrum coated laterally by a brush of stiff, spine-like setae and long fine setae, dorsal surface sparsely covered by long fine setae, anterior margin glabrous. Antennae elongate, with flagellomeres thin and long. Ocelli and compound eyes large, the latter reniform., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on page 4, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072

Key to genera and species of Haidomyrmecinae 1. Clypeus posteriorly with a slightly elevated setose lobe positioned just ventrad or between antennal insertion ������������������������������������������������������������������������.. ���������... 2 Clypeus posteriorly projecting forward into a cephalic horn arising ventrad, between, or dorsad antennal insertion ������������������������������������������������������������������������.. ���������... 5 2. Antenna compact, with flagellomeres short (������������������������������������������������������������������������������������ 3 Antenna elongate, with flagellomeres long (>2 as long as broad); first flagellomere shorter than second one ������������ ������������������������������������������������������ Haidomyrmex species: 4 3. All flagellomeres nearly of same length; clypeal setose pad positioned anterior to antennal insertion; ocelli present; two metatibial spurs ������������������������������������������������������������������ ��������������������� Haidomyrmodes mammuthus Perrichot et al., 2008. Flagellomere I longest of basal four flagellar articles; clypeal setose pad positioned slightly posterior to antennal insertion; ocelli absent; one metatibial spur ��������������������������� ���.. ���.. ������ Haidoterminus cippus McKellar, Glasier & Engel, 2013. 4. Flagellomere II longest of basal three flagellar articles; frontal triangle moderately elevated; vertex, ventral margin of mandibles, and antennomeres densely setose; mandibles with medioventral blade bearing at least one large apical tooth, with tip of apical portion tapered to acute point, smooth; ocelli reduced to pit-like traces; subpetiolar process present ���������������������������������. ��� Haidomyrmex cerberus Dlussky,1996 Flagellomere II longest; frontal triangle strongly elevated; vertex, ventral margin of mandibles, and antennomeres mostly glabrous; mandibles with medioventral blades asymmetrical, left one bearing three small teeth, right one bearing two larger teeth; tip of apical portion tapered to sharp point, smooth; ocelli and subpetiolar process absent ��������������������������������������� ������������������ Haidomyrmex scimitarus Barden & Grimaldi, 2012 Apical flagellomere longest; frontal triangle feebly elevated; vertex, ventral margin of mandibles, and antennomeres mostly glabrous; mandibles with medioventral blade bearing a single apical tooth, with tip of apical portion tapered to blunt point and minutely serrate; ocelli absent; subpetiolar process a minute tooth ������������������������������������������������������������������������������. ���������������������. Haidomyrmex zigrasi.. Barden & Grimaldi, 2012. 5. Posterior clypeal projection not furrowed ventrally (�� without lateral ridges), forming a pointed or paddle-shaped horn; long, paired trigger setae present, arising from base of clypeal process or ventral surface of apical setose pad; peg-like denticles absent or present only on ventral setose pad; labrum not obviously exposed ���������������������������������������������������������������������. ��� 6 Posterior clypeal projection distinctly furrowed ventrally, with lateral ridges originating above mandible insertions, and converging posteriorly to form the lateral margins of a horn; paired trigger setae absent; peg-like denticles present along lateral clypeal ridges, or on labrum, or both; labrum exposed ventrally of horn ������������������������������������������������������������ 9 6. Clypeal horn a small triangulate cuticular elevation; one pair of trigger setae positioned at base of horn; compound eyes small (OI 24); ocelli absent; flagellomeres IIeX compact, barely longer than wide ������������������������������������������������������������������������ ��������������������������� Protoceratomyrmex revelatus gen. et. sp. nov. Clypeal horn paddle-shaped, comprising basal stalk and apical setose pad; compound eyes small to large (OI 26e39); ocelli present; flagellomeres IIeX elongate, distinctly longer than wide ������������������������������������������������������������������������.... ��������� 7 7. Clypeal horn and curved apical portion of mandibles extremely long, reaching high above vertex (HoI 140; MDI 130); peg-like denticles present on ventral setose pad; 2 pairs of trigger setae positioned at base of horn's stalk; frontal triangle fused with horn, not distinguishable; gastral constriction between AIII and AIV at most faintly impressed ������������������������������������������������������������������������������������... ��������� Ceratomyrmex ellenbergeri Perrichot, Wang & Engel, 2016 Clypeal horn and curved apical portion of mandibles distinctly shorter, not reaching above vertex (MDI 70); peg-like denticles absent; 1 pair of trigger setae positioned on ventral surface of apical setose pad; frontal triangle connecting with horn by cleared cuticular membrane; gastral constriction between AIII and AIV deeply impressed ������������������������������������������������ ���������������������������������������������������. Linguamyrmex species: 8 8 Clypeal horn reduced (HoI 25e35), with stalk almost indistinct and apical setose pad hexagonal, as broad as (worker) or smaller than (gyne) compound eye's length (EL); mandibles with medioventral blade bearing two teeth and, in worker, surpassing midlength of apical curved portion; compound eyes small in worker (OI 25); propodeum higher than long ���.. ������������������. ������ Linguamyrmex brevicornis gen. et. sp. nov. Clypeal horn conspicuous (HoI 56), with stalk short, not surpassing frontal triangle in dorsal view, and apical setose pad circular, broader than EL in worker; mandibles with medioventral blade smooth and not reaching to midlength of apical curved portion; compound eyes large in worker (OI 39); propodeum longer than high ������������������������������������ ���������������������. ��� Linguamyrmex vladi Barden & Grimaldi, 2017. Clypeal horn enormous (HoI 95e105), with apical setose pad circular, nearly as broad as head in dorsal view, and stalk about as long as pad; mandibles with medioventral blade smooth and, in worker, not reaching to midlength of apical curved portion; compound eyes large in worker (OI 44); propodeum longer than high ������������������������������������������������������������������������������ ���������������������.. Linguamyrmex rhinocerus Miao & Wang, 2019. 9 Body and legs robust; mandibles scythe-shaped, with medioventral triangular blade present; in frontal view, the outline of apical portion of mandibles aligned with clypeal lateral ridges; clypeal horn gradually tapering to rounded tip; genae elongate, the mandibles inserted far from compound eyes ������������������������������������������������������������������������������.. ��� 10 Body and legs gracile; mandibles sickle-shaped, with medioventral triangular blade absent; clypeal horn and labrum protruding horizontally, their tip with a deep median notch; apical lateral margins of horn bilobed; genae strongly reduced, the mandibles inserted close to compound eyes ������������������������. ���������. Aquilomyrmex huangi gen. et. sp. nov. 10 Mandibles with apical curved portion much longer than basal portion and entirely serrate along dorsal margin; with medioventral blade an isosceles tooth not projecting beyond mandibular elbow (labrum and clypeal surface below horn entirely exposed when mandibles closed); ocelli large, antennomeres and compound eyes distinctly elongate ��������������������������������� Dhagnathos autokrator gen. et. sp. nov. Mandibles with apical curved portion not distinctly longer than basal portion; with medioventral blade a large triangle gradually tapering to tip of apical portion (labrum and clypeal surface below horn entirely concealed when mandibles closed); dorsal margin of mandibles serrate along distal half of medioventral projection, bidentate apically; ocelli, antennomeres, and compound eyes more reduced ������������ ������������������������������������ ������������������������������������������ Chonidris insolita gen. et. sp. nov., Published as part of Perrichot, Vincent, Wang, Bo & Barden, Phillip, 2020, New remarkable hell ants (Formicidae: Haidomyrmecinae stat. nov.) from mid-Cretaceous amber of northern Myanmar, pp. 1-18 in Cretaceous Research 109 on pages 16-17, DOI: 10.1016/j.cretres.2020.104381, http://zenodo.org/record/3653072, {"references":["Perrichot, V., Lacau, S., Neraudeau, D., Nel, A., 2008 b. Fossil evidence for the early ant evolution. Naturwissenschaften 95, 85 e 90. https: // doi. org / 10.1007 / s 00114 - 007 - 0301 - 8.","McKellar, R. C., Glasier, J. R. N., Engel, M. S., 2013 a. A new trap-jawed ant (Hymenoptera: Formicidae: Haidomyrmecini) from Canadian Late Cretaceous amber. The Canadian Entomologist 145, 454 e 465. https: // doi. org / 10.4039 / tce. 2013.23.","Dlussky, G. M., 1996. Ants (Hymenoptera: Formicidae) from Burmese amber. Paleontological Journal 30, 449 e 454. Translated from Paleontologicheskii Zhurnal 1996 (3), 83 e 89.","Barden, P., Grimaldi, D. A., 2012. Rediscovery of the bizarre Cretaceous ant Haidomyrmex Dlussky (Hymenoptera: Formicidae), with two new species. American Museum Novitates 3755, 1 e 16. https: // doi. org / 10.1206 / 3755.2.","Perrichot, V., Wang, B., Engel, M. S., 2016. Extreme morphogenesis and ecological specialization among Cretaceous basal ants. Current Biology 26, 1468 e 1472. https: // doi. org / 10.1016 / j. cub. 2016.03.075.","Barden, P., Herhold, H. W., Grimaldi, D. A., 2017. A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure. Systematic Entomology 42, 837 e 846. https: // doi. org / 10.1111 / syen. 12253.","Miao, Z., Wang, M., 2019. A new species of hell ants (Hymenoptera: Formicidae: Haidomyrmecini) from the Cretaceous Burmese amber. Journal of Guangxi Normal University 37 (2), 139 e 142. https: // doi. org / 10.16088 / j. issn. 1001 - 6600.2019. 02.017."]}

Lioponera neocaledonica Jouault, Ramage & Perrichot sp. nov. urn:lsid:zoobank.org:act: 7612551C-6C0A-4931-9D06-8FC8062DB5E3 Figs 4, 5B Diagnosis Lioponera neocaledonica Jouault, Ramage & Perrichot sp. nov. is mostly characterized by the combination of the following features: mesosoma dorsolaterally conspicuously marginate, propodeal declivity without dorsal margin; posterolateral corners of petiole not projected into spines. The two latter features easily serve to distinguish L. neocaledonica Jouault, Ramage & Perrichot sp. nov. from the other New Caledonian species L. cohici (Wilson, 1957) (Fig. 5A) and L. dumbletoni (Wilson, 1957) (Fig. 5C). Etymology The specific epithet refers to New Caledonia where the species was discovered. Type material Holotype NEW CALEDONIA ��� wingless ♀; Grande Terre, Province Sud, Sarram��a, Plateau de Dogny trail; 21��37���31��� S, 165��52���5��� E; 441 m; 16 Nov. 2011; V. Perrichot leg. [VP188]; mixed mountain forest, hand picking on ground; MNHN. Paratypes NEW CALEDONIA ��� 2 wingless ♀♀; same collection data as for holotype; MNHN ��� 1 wingless ♀; Grande Terre, Province Sud, Montagne des Sources; 22��09���35.1��� S, 166��35���42.4��� E; 394 m; 23 Oct. 2010; M. Berman leg. [CSIRONC0055]; ANIC 32-146088. Description (holotype) MEASUREMENTS (in mm; holotype in brackets). TL 4.12���4.15 (4.15); HL 0.8���0.85 (0.85); HW 0.7���0.77 (0.77); SL 0.45���0.52 (0.52); ED 0.2���0.25 (0.2); PW 0.65���0.72 (0.7); MH 0.52���0.62 (0.62); MSL 1.025��� 1.12 (1.12); PL 0.45���0.55 (0.55); PH 0.4���0.42 (0.42); DPW 0.67���0.7 (0.7); T1W 0.65���0.77 (0.77); T1L 0.5���0.62 (0.62); T2W 0.75���0.82 (0.82); T2L 0.6���0.65 (0.65). Indices: CI 84���91 (91); SI 64���69 (67); LPI 76���94 (77); DPI 127���150 (127). HEAD (Fig. 4C). Sub-rectangular, slightly longer than broad, widest above midline of head; lateral sides slightly convex; vertexal margin slightly concave, posterior lateral corners slightly rounded. Parafrontal ridges angular. Compound eyes of medium size, situated below midline of head. Mandible elongate, triangular, masticatory margin with 10 small teeth. Antennae with scape short and clavate, reaching up to two fifth of posterior margin of head. MESOSOMA (Fig. 4 A���B, D). Stout, roughly rectangular in dorsal view, with conspicuous dorsolateral margins, the margins notched around promesonotal junction; mesosomal dorsal surface convex, propodeal declivity without dorsal margin. METASOMA. Petiole (Fig. 4 A���B, E) broader than long, slightly trapezoidal, widest at two thirds of length; anterior petiolar surface concave; dorsal petiolar surface convex, with dorsolateral margins well developed and convex, without anterodorsal and posterodorsal margins; posterior petiolar surface slightly convex; anterodorsal and posterodorsal corners angular. Subpetiolar process small, triangular, with stout acute somewhat hooked apex; fenestra absent. Gaster elongate (Fig. 4 A���B), with distinct girdling constriction (cinctus) between abdominal segments III and IV.Abdominal segment II trapezoidal, slightly broader than long, widest at one third of length; dorsal surface slightly convex; dorsolateral margins well developed and strongly convex; anterodorsal margin sinuate; anterodorsal corner broadly rounded; posterolateral corner angular. Sting exerted, upcurved. INTEGUMENT. Mostly black except last two funicular articles, tarsomeres, and posterior margins of abdominal segments IV to V brown. Body finely and sparsely punctate; reticulation present on sides of the mesosoma, becoming shallower from propleura to propodeum; cinctus, abdominal segment IV, base of abdominal segments V, VI and pygidium transversally, finely and shallowly striate. Body covered with suberect hairs; scape with abundant appressed hairs; tibiae with scattered decumbent hairs; tarsi with abundant decumbent to appressed hairs; declivous face of propodeum with short, dense and appressed hairs. Remarks The potential presence of ergatogynes in Lioponera (Wilson 1957; Borowiec 2016) prevents assignment of the present specimens to workers or ergatogynes. Key to the species of Lioponera of New Caledonia (modified from Wilson 1957) 1. Large species (HW> 1.1 mm), with long scapes reaching nearly to posterior corners of head; mesosomal dorsolateral margins absent................................ Lioponera dumbletoni (Wilson, 1957) ��� Small species (HW Lioponera cohici (Wilson, 1957) ��� Mesosomal posterodorsal margin absent (Fig. 4D); posterolateral corners of petiole not projected into spines (Fig. 5B)..................... Lioponera neocaledonica Jouault, Ramage & Perrichot sp. nov., Published as part of Ramage, Thibault, Jouault, Corentin, Schmidt, Alexander R., Seyfullah, Leyla J. & Perrichot, Vincent, 2019, Two new ant species (Formicidae: Dorylinae, Ponerinae) from New Caledonia, pp. 1-14 in European Journal of Taxonomy 589 on pages 7-8, DOI: 10.5852/ejt.2019.589, http://zenodo.org/record/3659941, {"references":["Wilson E. O. 1957. The discovery of cerapachyine ants on New Caledonia, with the description of new species of Phyracaces and Sphinctomyrmex. Breviora 74: 1 - 9.","Emery C. 1914. Les fourmis de la Nouvelle-Caledonie et des iles Loyalty. Nova Caledonia. Forschungen in Neu-Caledonien und auf den Loyalty-Inseln. Recherches scientifiques en Nouvelle-Caledonie et aux iles Loyalty. A. Zoologie 1: 393 - 437. https: // doi. org / 10.5962 / bhl. title. 6790","Wilson E. O. 1958. Studies on the ant fauna of Melanesia. I. The tribe Leptogenyini. II. The tribes Amblyoponini and Platythyreini. Bulletin of the Museum of Comparative Zoology 118: 101 - 153.","Borowiec M. L. 2016. Generic revision of the ant subfamily Dorylinae (Hymenoptera, Formicidae). ZooKeys 608: 1 - 280. https: // doi. org / 10.3897 / zookeys. 608.9427"]}

Leptogenys loarelae Ramage sp. nov. urn:lsid:zoobank.org:act: 819E8C18-56F7-4CB5-AE07-00B5406D924A Fig. 1 Diagnosis Leptogenys loarelae Ramage sp. nov. can be distinguished from other New Caledonia species of Leptogenys by the unique combination of the following features: integument smooth and shining, mostly black colored with blueish reflection; third antennal segment markedly elongate, more than twice the length of pedicel; mandible elongate, triangular; petiole elongate, nearly three times longer than broad in dorsal view. Etymology The specific epithet refers to the first name of Thibault Ramage���s daughter, Loarel. The species epithet is to be treated as a noun in the genitive case. Type material Holotype NEW CALEDONIA ��� wingless ♀; Grande Terre, Province Sud, Sarram��a, Plateau de Dogny trail; 21��37���10.7��� S, 165��52���31.9��� E; 897 m; 16 Nov. 2011; V. Perrichot leg. [VP209]; mixed mountain forest, hand picking on ground; MNHN. Description Worker holotype MEASUREMENTS (in mm). TL 11.5; HL 2.06; HW 1.3; SL 2.56; ED 0.5; PW 1.16; MSL 3.65; PL 1.83; PH 0.8; DPW 0.63; CI 62.9; SI 197.4; LPI 43.6; DPI 34.5. HEAD. Elongate, sub-trapezoidal, lateral sides diverging anteriorly (Fig. 1B); lateral margin barely convex; posterior margin weakly concave medially, posterior lateral corners rounded. Clypeus triangular, with blunt apex and sligthly concave side margins. In full-face view, eye large, oval, maximum diameter greater than twice the maximum width of scape; placed at head midline. Antennae with scape long and slightly enlarged medially, surpassing posterior margin of head by its 2������ th; funicular segments elongate; third antennal segment markedly elongate, more than twice the length of the second. Mandible triangular, elongate, shorter than head width; masticatory margin edentate. MESOSOMA (Fig. 1A, C). Elongate, compressed laterally, widest at pronotum; mesosomal dorsal surface in profile slightly convex. In dorsal view, mesonotum as long as broad. Propodeal spiracle slit-like. Petiole (Fig. 1A, C���D) elongate, nearly three times as long as broad, trapezoidal, widest at apex; dorsal surface of petiolar node convex, posterior surface slightly concave, anterodorsal and posterodorsal corners angular. Subpetiolar process small, trapezoidal; located beneath anterior 1������ th of the petiole (Fig. 1D). Gaster elongate. INTEGUMENT. Smooth and shining, very finely and sparsely punctate; mostly black with blueish reflection, except mandibles, antennal radicel and funicular articles, tarsomeres and hypopygium and pygidium reddish brown. Dorsum of body covered by erect and suberect yellow setae; setae longer on head, scape, pronotum, procoxa and gaster, but shorter on remaining body. Key to the Leptogenys species (workers) of New Caledonia (modified from Wilson 1958) 1. Large species (HW> 1.2 mm).......................................................................................................... 2 ��� Small species (HW ��� 1.2 mm).......................................................................................................... 3 2. Mandibles exceptionally long, their length measured from insertion to tip exceeding the head width (Fig. 2A)............................................................................................ Leptogenys rouxi Emery, 1914 ��� Mandibles shorter, their length much less than the head width (Fig. 1B)................................................................................................................................. Leptogenys loarelae Ramage sp. nov. 3. Head entirely covered by coarse punctures (Fig. 2C).................. Leptogenys punctata Emery, 1914 ��� Head entirely smooth and shining (Fig. 2B)..................................................................................... 4 4. Larger species, head width ��� 0.9 mm; in dorsal view, posterior corners of petiolar node forming distinct angles of 45���50�� (Fig. 3A).......................................... Leptogenys acutangula Emery, 1914 ��� Smaller species, head width not exceeding 0.75 mm; in dorsal view, posterior corners of petiolar node thick and broadly rounded (Fig. 3B)..................................... Leptogenys sagaris Wilson, 1958 Subfamily Dorylinae Leach, 1815 Genus Lioponera Mayr, 1879, Published as part of Ramage, Thibault, Jouault, Corentin, Schmidt, Alexander R., Seyfullah, Leyla J. & Perrichot, Vincent, 2019, Two new ant species (Formicidae: Dorylinae, Ponerinae) from New Caledonia, pp. 1-14 in European Journal of Taxonomy 589 on pages 4-7, DOI: 10.5852/ejt.2019.589, http://zenodo.org/record/3659941, {"references":["Wilson E. O. 1958. Studies on the ant fauna of Melanesia. I. The tribe Leptogenyini. II. The tribes Amblyoponini and Platythyreini. Bulletin of the Museum of Comparative Zoology 118: 101 - 153.","Emery C. 1914. Les fourmis de la Nouvelle-Caledonie et des iles Loyalty. Nova Caledonia. Forschungen in Neu-Caledonien und auf den Loyalty-Inseln. Recherches scientifiques en Nouvelle-Caledonie et aux iles Loyalty. A. Zoologie 1: 393 - 437. https: // doi. org / 10.5962 / bhl. title. 6790"]}

Two new species of ants (Hymenoptera: Formicidae) collected from New Caledonia are described and figured based on worker specimens: Leptogenys loarelae Ramage sp. nov. (Ponerinae, Ponerini) and Lioponera neocaledonica Jouault, Ramage & Perrichot sp. nov. (Dorylinae, Cerapachyini). All specimens were collected from the South Province of Grande Terre. These two new species are primarily distinguished from the other New Caledonian relatives by the size and shape of petiole for L. loarelae Ramage sp. nov. and by the presence of dorsolateral margins on the mesosoma for L. neocaledonica Jouault, Ramage & Perrichot sp. nov. Keys to New Caledonian Leptogenys and Lioponera are provided.

International audience; Examination of fossil ant specimens from various private and institutional German collections of Baltic amber has revealed a new genus and species of Myrmicinae, described here as Thanacomyrmex hoffeinsorum gen. et sp. nov. The new worker morphotype shares some similarities with the coeval extinct genus Parameranoplus Wheeler, also from Baltic amber, and the extant genera Pristomyrmex Mayr and Acanthomyrmex Emery, suggesting placement of the new fossil in the Myrmecina genus-group within the tribe Crematogastrini. The new genus highlights the tremendous diversity of ants that evolved in the Palaeogene European forests and composed the first major radiation of crown-group ants.; L’examen de fourmis fossiles dans diverses collections privées et institutionnelles allemandes d’ambre balte a révélé un nouveau genre et une nouvelle espèce de Myrmicinae, décrits ici comme Thanacomyrmex hoffeinsorum gen. et sp. nov. Ce nouveau morphotype d’ouvrière partage des similitudes avec le genre éteint Parameranoplus Wheeler, également connu de l’ambre balte, et les genres actuels Pristomyrmex Mayr et Acanthomyrmex Emery, suggérant le placement du nouveau fossile dans le groupe de genres Myrmecina, parmi la tribu Crematogastrini. Le nouveau genre souligne la grande diversité de fourmis ayant évolué dans les forêts paléogènes d’Europe et constituant la première radiation majeure du groupe couronne des fourmis.