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Ilyocoris andancensis sp. nov. (Figs 1–4) urn:lsid:zoobank.org:act: 3EB380F0-1632-4971-A5ED-7DFE8A2DC9FA Material. Holotype MNHN-F.A71364 (Romain Jattiot coll.), paratypes MNHN-F.A71365, and MNHNF.A71366 (all compressed mummies), stored at the Muséum National d’Histoire Naturelle, Paris, France. Diagnosis. Adult characters only. Color patterns identical on the three fossils. Connexivum brownish along posterior part and yellowish along anterior part; green-yellow anterior part of pronotum clearly narrower than black posterior part; outer margin of embolium black. Type locality and horizon. Lower limit of Turolian to Tortonian-Messinian boundary, 7.30 ± 0.15 Ma (Pastre et al., 2004), ‘ La Montagne d’Andance’, Saint-Bauzile village, Ardèche, France. Description (based on all specimens). Head transverse; ocelli absent; eyes sessile; labium short and thick, arising near apex of head, conical, obviously segmented, without transverse sulcation; labium; apex of head without a laminated extension; vertex less than two times and half broader than transversal diameter of compound eye; head, pronotum, and scutellum shiny; ground color black and green-yellow; brown dots on head, some of which form two longitudinal, parallel, median lines, each of them widening posteriorly; anterior margin of pronotum straight behind interocular space; lateral pronotal margins limited inside by a strong sunken line; anterior part of pronotum green-yellow, and posterior part black; scutellum black; no meso- and metasternal midventral carinae (observed on holotype by partly removing the mummy); macropterous, hemelytra olive brown, opaque, very finely and densely punctate, clavus and margin paler, embolium with a less dense punctuation than the rest of hemelytra but with outer margin black; membrane without venation; fore legs raptorial, fore femur only moderately incrassate with inner antero-basal angle not produced, forming a continuous curve with trochanter, tarsus without a fringe of setae; hind legs flattened; tibiae with strong brownish spines and long swimming setae. Legs pale yellow; apex of abdomen without respiratory processes; connexivum brownish along posterior part and yellowish along anterior part. Dimensions. Body 14.7 mm long, 6.4 mm wide (MNHN-F.A71364), 12.0 mm long, 6.0 mm wide (MNHN-F.A71365), 13.5 mm long, 6.0 mm wide (MNHN-F.A71366); head 1.3 mm long, 3.4 mm wide (MNHN-F.A71364); pronotum 2.0 mm long, 4.1 mm wide (MNHN-F.A71364); abdomen 9.7 mm long, 6.5 mm wide (MNHN-F.A71364); fore tibia 2.1 mm long, 0.9 mm wide; hind tibia 3.0 mm long, tarsus 4.1 mm long, hemelytra 8.6 mm long (MNHN-F.A71364).

Genus Ilyocoris Stål, 1861 Type species. Nepa cimicoides Linnaeus, 1758; by original designation., Published as part of JATTIOT, ROMAIN, TRINCAL, VINCENT, GARROUSTE, ROMAIN & NEL, ANDRÉ, 2023, A new fossil species of the saucer bug genus Ilyocoris (Heteroptera, Naucoridae) from the Upper Miocene maar paleolake of la Montagne d'Andance (France), pp. 260-267 in Palaeoentomology 6 (3) on page 261, DOI: 10.11646/palaeoentomology.6.3.9, http://zenodo.org/record/8073120

JATTIOT, ROMAIN, TRINCAL, VINCENT, GARROUSTE, ROMAIN, NEL, ANDRÉ (2023): A new fossil species of the saucer bug genus Ilyocoris (Heteroptera, Naucoridae) from the Upper Miocene maar paleolake of la Montagne d'Andance (France). Palaeoentomology 6 (3): 260-267, DOI: 10.11646/palaeoentomology.6.3.9, URL: http://dx.doi.org/10.11646/palaeoentomology.6.3.9

Eornithoica grimaldii sp. nov. (Fig. 1) urn:lsid:zoobank.org:act: 6234ECDE-4652-4FA1-9A6D- F765A993D8D8 Holotype. FOBU17740 (part and counterpart of a complete fly), deposited in the collection of Fossil Butte National Monument, Wyoming, USA. Etymology. The specific epithet is dedicated to Prof. Dr David A. Grimaldi, in recognition of his impressive contributions to the study of fossil insects and extant Diptera. Diagnosis. As for the genus (vide supra). Type locality and horizon. Douglass Pass, Garfield County, Colorado, USA, Early Eocene, Green River Formation, Parachute Creek Member, Ypresian, 46.2– 40.4 Ma. Description. Body ca. 7.0 mm long, as preserved, dark brown, covered with numerous stout spines and spurs; head poorly preserved, detached from thorax but originally deeply inserted between fore legs, ca. 1.2 mm long, 1.0 mm wide, with mouthparts projected forward; compound eyes in lateral positions; ocelli not visible, apparently absent. Thorax 3.1 mm long, 2.1 mm wide; thoracic segmental borders indistinct dorsally, but not completely obliterated; procoxae widely separated, with a large, distinctly less-sclerotized area between them; protergum distinctly shortened; mesothorax wider than long; mesocoxae widely separated; metacoxae widely separated; wings normally developed, 6.4 mm long, ca. 2.3 mm wide, slightly shorter than body; R1 short; R2+3 longer than R1, reaching wing midlength; R4+5, M1+2, and M3+4 long; crossveins r-m and i-m present but faint; R4+5 reaching wing margin distally with a broad cell between R2+3 and R4+5; eight long and stout spurs on costal margin; numerous strong spines/spurs on legs; procoxa inflated, greatly enlarged; tarsomeres I longer than tarsomeres II to IV, the latter distinctly shortened and compact; pretarsal claws recurved, not bifid, with a conspicuous base or heel; at least apex of mesotibia with a row of five stout spurs; mesofemur stout, as long as mesotibia; abdomen 3.0 mm long, 2.9 mm wide, covered with numerous long, stout spines/spurs, some situated on enlarged sclerotized bases; three large transverse dorsal sclerotized plates on basal-most segments. Remarks. It is impossible to determine what host might have been visited by the present fossil, but given that basal hippoboscids are typically ectoparasitic on birds, it is likely that E. grimaldii was specialized for some extinct species of Aves. The bird fauna of the Green River Formation is rather diverse and certainly harbored any number of potential hosts for these flies (Weidig, 2010), but fossil bats are also present in this formation (Jepsen, 1966).

Eornithoica gen. nov. urn:lsid:zoobank.org:act: 0C3395B4-3ED4-49D3-BA57- 0F65525B4C16 Type species. Eornithoica grimaldii sp. nov. Etymology. The new generic name is taken from the Ancient Greek goddess of the dawn, Ēṓs (Ἠώς, and from which the Eocene epoch derives its name), the genus name Ornithoica. The gender of the name is feminine. Diagnosis. Body, especially abdomen, and legs with numerous stout spines; dorsal segmentation of thorax not completely obliterated; wings fully developed; eight long and stout spurs on costal margin of wing; R1 short, R4+5 reaching wing margin distally, with a broad cell between R2+3 and R4+5; pretarsal claws with weakly developed basal heels; abdomen with three large transverse dorsal sclerotized plates., Published as part of NEL, ANDRÉ, GARROUSTE, ROMAIN & ENGEL, MICHAEL S., 2023, The earliest Pupipara (Diptera: Hippoboscoidea): A new genus and species from the lower Eocene of the Green River Formation, pp. 58-63 in Palaeoentomology 6 (1) on page 59, DOI: 10.11646/palaeoentomology.6.1.9, http://zenodo.org/record/7754965

Lodevoisadia coheni gen. et sp. nov. urn:lsid:zoobank.org:act: 59BE288B-C752-4898-ACE5-72799793A9F6 Fig. 1 Diagnosis As for the genus (vide supra). Etymology Named after Henri Cohen, who discovered the type specimen and allowed us to study it. The specific epithet is to be treated as a noun in genitive case. Type material Holotype Specimen 2022.3.2 (number HC Can 01a-b, part and counterpart of a forewing, collection Henri Cohen); Musée of Lodève, France. Description Counterpart of an almost complete right forewing, with costal part of extreme base not preserved, 10.4 mm long (as preserved), ca 3.5 mm wide; area between C and ScP narrow, ca 0.2 mm wide; ScP simple, closely parallel to R, ending in C at 8.0 mm from wing base; few weak and short veinlets between RA and C near apex of RA; base of RP at 3.3 mm from wing base; RP simple; base of MP at 2.5 mm from wing base; MA simple; MP forked 3.3 mm distad origin of MP; crossveins in areas between RA and MP spaced; course of RP and MA nearly straight; areas between RA and RP and between RP and MA distally widened; CuA convex, with a nearly straight and simple basal posterior branch CuA2; CuA1 branched at 2.0 mm distad its base; area between MP and CuA1 rather broad basad first branch of CuA1; RP, M, and CuA well-separated; arculus m-cua present between M and CuA1; CuP concave, straight, and simple; AA1 and AA2 convex and simple.

Genus Lodevoisadia gen. nov. urn:lsid:zoobank.org:act: CB082E08-C310-4BBD-ACE8-FEFA3D7DC580 Type species Lodevoisadia coheni gen. et sp. nov. Diagnosis Costal area narrow; ScP ending into C; RP simple; MA simple; MP forked; CuA1 with three branches; CuA2 simple; arculus m-cua present between M and CuA1; areas between RA and RP, RP and MA, and between MA and MP distally widened; few simple crossveins between RP, MA, and MP. Etymology The genus name is a combination of the town of Lodève and the genus name Isadia. Gender feminine., Published as part of Nel, André, Kundura, Jean-Paul, Pouillon, Jean-Marc, Garrouste, Romain & Jouault, Corentin, 2022, A new ' grylloblattodean' genus and species (Insecta: Polyneoptera) from the middle Permian Salagou Formation (France), pp. 77-84 in European Journal of Taxonomy 852 (1) on page 79, DOI: 10.5852/ejt.2022.852.2011, http://zenodo.org/record/7459029

Eoacridophagus azari sp. nov. (Figs. 1���2) Material. Holotype MNHN.F. A 57338, stored in the Laboratory of Palaeontology, MNHN, Paris, France. Diagnosis. That of the genus. Very small fly, wing 1.3 mm long. Etymology. Named after our friend Dr Dany Azar (Lebanese University), specialist on fossil insects. Type horizon. Lowermost Eocene, Sparnacian, level MP 7 of the mammal fauna of Dormaal. Type locality. Le Quesnoy, Chevri��re, region of Creil, Oise department, France. Description. Head 0.3 mm long and 0.25 mm wide, 0.4 mm high; eyes large but dichoptic, well separated; three ocelli placed in triangle on posterior margin of vertex; antennae shorter than head, 0.2 mm long; flagellum broader in its basal third than at apex; proboscis shortened, not surpassing the mouthpart cavity. Thorax 0.45 mm long and 0.55 mm high; mesonotum covered with a short pilosity; lateral sides bare. Wing 1.3 mm long, 0.5 mm wide; cell r 1 not a small triangle distally opened; apex of vein R 1 0.7 mm from wing base; Rs 0.1 mm long; R 2 + 3 ending in costa well separated from R 1, 0.4 mm long; vein R 4 + 5 0.75 mm long, ending in costa at a level clearly well beyond end of vein M 2, not reaching wing apex; r-m at 0.55 mm from wing base, approximately in middle of wing; vein M 2 present; cell cup closed just before posterior wing margin. Legs long and slender, fore femur 0.4 mm long, tibia + tarsus 0.6 mm; mid leg 0.95 mm long; hind femur 0.45 mm long, tibia + tarsus 0.5 mm long; femora bare; tibia covered with short bristles; no tibial spur. Abdomen deformed, 0.6 mm long, 0.4 mm wide, covered with short hairs. Genitalia not clearly visible. Discussion. Following the key to families of Bombyliidae s.l. of Greathead & Evenhuis (2001), our fossil can be included in the Mythicomyiidae because of the following characters: wing with R 4 + 5 not branched; vein MA absent; palpi absent; abdominal spiracles located in terga; very small insect (wing 1.3 mm long). Eoacridophagus gen. nov. falls in the Psiloderoidinae because of the following characters: wing vein R 4 + 5 ending in costa at a level clearly well beyond end of vein M 2; R 2 + 3 present and ending in costa well separated from R 1; cell r 1 large, not a small triangle; antennal style placed apically on second flagellomere; vein R 2 + 3 short, much shorter than R 4 + 5. The Platypyginae have their vein R 2 + 3 long, similar in length to R 4 + 5, unlike Eoacridophagus. Following the key to the modern genera of Psiloderoidinae presented by Lamas et al. (2015), Eoacridophagus differs from Psiloderoides Hesse, 1967 for the proboscis probably functional even if it is very short. Acridophagus Evenhuis, 1983 (replacement name for Cyrtomorpha White, 1916) has a proboscis half then length of the head, and projecting forward, longer than in Eoacridophagus. Their wing venations are very similar (White, 1916). Amydrostylus Lamas, Falaschi & Evenhuis, 2015 has a very long proboscis but its venation is very similar to that of Eoacridophagus. Eoacridophagus also differs from Onchopelma Hesse, 1938 in the narrower anal area, cell cup closed before the wing margin by a stalk (Hesse, 1938; Evenhuis, 2002). Eoacridophagus differs from Psiloderoides in the absence of the discal cell, r-m situated well basal of level of apex of R 1, and R 2 + 3 longer (Hesse, 1967). In addition, our fossil differs from all the fossils of the Psiloderoidinae: the genera Proplatypygus Hennig, 1969 (Eocene Baltic amber) and Borissovia Evenhuis, 2002 (Cretaceous amber of Siberia) can easily be separated from Eoacridophagus by the different antennal shape, the presence of a discal cell and the well-developed mouthparts (Hennig, 1969; Zaitzev, 1986; Evenhuis, 2002). Carmenelectra Evenhuis, 2002 (Eocene Baltic amber) differs from Eoacridophagus in the well-developed proboscis (Evenhuis, 2002). Procyrtosia Zaitzev, 1986 (Cretaceous amber of Siberia) differs from Eoacridophagus in the presence of a discal cell, CuA 2 and A 1 not fused, and a long vein R 2 + 3. The proportions between the lengths of R 2 + 3 and R 4 + 5 in Procyrtosia are similar to the situation in Platypygus (Platypyginae), suggesting a position in this last subfamily. Eoacridophagus differs from Palaeoplatypygus Kovalev, 1985 (Cretaceous amber of Siberia) in the absence of discoidal cell, shorter R 1 and CuA 2 -A 1 fusion on the ventral margin of wing (Kovalev, 1985; Zaitzev, 1986). Zarzia Zaitzev, 1986 (Cretaceous amber of Siberia) has a vein R 2 + 3 sigmoidal, and a discal cell unlike our fossil. Remark. The recent Psiloderoidinae are known to be parasitoids on the eggs of Acridoidea. Grasshoppers are really not frequent in the Oise amber, which in no way means that they were not present in the corresponding palaeoenvironment because these insects are quite able to escape the embedment in the fresh resin. The recent Psiloderoidinae are distributed in the Neotropical, Afrotropical and Australian areas, in warm environments, which is in accordance with the current reconstruction of the palaeoenvironment of the Oise amber and the presence of other insects also living in such biota (viz. Isoptera Mastotermitidae) (Nel & Bourguet, 2006; Nel & Brasero, 2010)., Published as part of Myskowiak, Justine, Garrouste, Romain & Nel, Andre, 2016, A new genus and species of micro bee fly from the Earliest Eocene French amber (Diptera: Mythicomyiidae: Psiloderoidinae), pp. 583-586 in Zootaxa 4114 (5) on pages 583-586, DOI: 10.11646/zootaxa.4114.5.5, http://zenodo.org/record/259075, {"references":["Greathead, D. J. & Evenhuis, N. L. (2001) Annotated keys to the genera of African Bombylioidea (Diptera: Bombyliidae; Mythicomyiidae). African Invertebrates, 42, 105 - 224.","Lamas, C. J. E., Falaschi, R. L. & Evenhuis, N. L. (2015) A new genus and species of micro bee flies from Brazil (Diptera: Mythicomyiidae: Psiloderoidinae). Zootaxa, 3955 (3), 403 - 416. http: // dx. doi. org / 10.11646 / zootaxa. 3955.3.7","Hesse, A. J. (1967) Additions to the Cyrtosiinae (Bombyliidae) of South Africa. Annals of the South African Museum, 50, 89 - 130.","White, A. (1916) The Diptera Brachycera of Tasmania. Part 3, families, Asilidae, Bombyliidae, Empidae, Dolichopodidac and Phoridae, Pap. Proceedings of the Royal Society of Tasmania, 1916, 148 - 266. http: // dx. doi. org / 10.5962 / bhl. part. 28993","Hesse, A. J. (1938) A revision of the Bombyliidae (Diptera) of southern Africa. Part 1. Annals of the South African Museum, 34, 1 - 1053.","Evenhuis, N. L. (2002) Review of the Tertiary microbombyliids (Diptera: Mythicomyiidae) in Baltic, Bitterfeld, and Dominican amber. Zootaxa, 100, 1 - 15.","Hennig, W. (1969) Kritische Betrachtungen uber der phylogenetischen Bedeutung von Bernsteinfossilien: die Gattungen Proplatypygus (Diptera, Bombyliidae) und Palaeopsylla (Siphonaptera). Memorie della Societa Entomologica Italiana, 48, 57 - 67.","Zaitzev, V. F. (1986) New species of Cretaceous fossil bee flies and a review of paleontological data on Bombyliidae (Diptera). Entomologicheskoe Obozrenie, 65, 815 - 825. [English translation in 1987. Entomological Review, 66, 150 - 160]","Kovalev, V. G. (1985) [Infraorder Tabanoidea Leach, 1815.] In: Kalugina, N. S. & Kovalev, V. G. (Eds.), Dvukrylye Yury Sibiri [Insecta Diptera of the Jurassic in Siberia.] Akademiya nauk SSSR, Paleontologicheskij Instituta, Moskovskogo Obshchestva Ispytatelej Prirody, pp. 183 - 187. [in Russian]","Nel, A. & Bourguet, E. (2006) Termite of the Early Eocene amber of France (Isoptera: Mastotermitidae, Kalotermitidae). Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte, 2006 (2), 101 - 115.","Nel, A. & Brasero, N. (2010) Oise amber. In: Penney, D. (Ed.), Biodiversity of fossils in amber from the major world deposits. Siri Scientific Press, Manchester, pp. 137 - 148."]}

Myskowiak, Justine, Garrouste, Romain, Nel, Andre (2016): A new genus and species of micro bee fly from the Earliest Eocene French amber (Diptera: Mythicomyiidae: Psiloderoidinae). Zootaxa 4114 (5): 583-586, DOI: http://doi.org/10.11646/zootaxa.4114.5.5

Hypopsylla belmontensis sp. nov. (Figs 1, 2) Material. Holotype In 45397/ In 46044 (print and counterprint) (The Natural Museum, London). Age and outcrop. Late Permian, mid Lopingian, cca 255 Mya, Newcastle Coal Measures, Warners Bay - Belmont area near Newcastle, New South Wales, Australia (Shi et al. 2010). Etymology. Named after the type locality Belmont. Description. Print and counterprint of two wings partly overlapping and deformed, these wings being the fore- and the hind wings due to their relative positions. Their venations are nearly identical. Forewing nearly complete, fragment 6.0 mm long, 2.5 mm wide in widest part; ScP ending on RA about 2.7 mm from wing base, but with a crossvein between it and costa; area between radius and costa broad, 0.25 mm wide; R, M and CuA fused into a common stem at wing base, making a weak posterior curve for about 1.1 mm; then M+CuA and radius separating, with R and basal stem R+M+CuA making a pronounced angle at this point; RP and RA separating 1.2 mm distal of base of M+CuA; convex RA with a deep posterior curve limiting a dark pigmented pterostigma, at least 1.2 mm long and 0.5 mm wide, pterostigma limited by a vein basally (ScP); a crossvein perpendicular to RA and to RP below middle of pterostigma; relatively concave RP without visible fork in preserved part; M and CuA separating immediately distal of point of reemergence of M+CuA; neutral stem of M very long, 2.5 mm long before its first fork; anterior branch of M distally with a deep fork; posterior branch of M with three branches ending on posterior wing margin; convex CuA very short before crossvein cua-cup ends on its, being aligned with distal part of CuA; distal part of CuA very long, 2.5 mm long before areola postica; areola postica very long and narrow, parallel to posterior wing margin, with CuA1 curved and CuA2 very short; cua-cup weaker than CuA and M, 0.9 mm long from base of CuP and CuA; concave CuP weakly curved and simple; two convex simple anal veins basally curved; anal area about 2.2 mm long along posterior wing margin. Discussion. The pattern of wing venation of Hypopsylla is typical of the Acercaria with a common stem R+M+CuA and a neutral crossvein cua-cup between concave CuP and convex CuA weaker than CuA (Nel et al. 2012). Hypopsylla has the wing venation synapomorphies of the Permopsocida Tillyard, 1926, as redefined in Huang et al. (2016), viz. fore- and hind wings with sclerotized pterostigma posteriorly limited by RA, with RA making a strong angle at the point of contact with crossvein between it and RP (synapomorphy of the Permopsocida, a structure found also in some Psocodean forewing but never in their hind wing); a distinct angle of radial vein at base of M+CuA (a character absent in Psocodea and Thripida, but present to a certain point in the forewings of many Hemiptera of different lineages, viz. the Archescytinidae, Prosbolidae, Pincombeidae, etc.); presence of an areola postica in forewings (probably an acercarian symplesiomorphy as this structure is present in Psocodea and Hemiptera but reduced in Thripida); presence of an areola postica (forked CuA) in hindwings, a character present in Hemiptera but very rare in Psocodea (in some specimens of Epipsocus lucifugus (Rambur, 1842)), and absent in the Thripida; RP with a simple apical fork (structure also present in Psocodea, Thripida, and Hemiptera); M with two main branches, each forked again (a character present in Hemiptera, Thripida have either a simple or a forked M, while Psocodea have a three- to simple M). The ScP either parallel to radius, or fused to costal margin, re-emerging from it or not, are character states present in various Psocoptera. Hypopsylla differs from the other representatives of the clade Permopsocida in the much deeper posterior curve of RA in the pterostigma and the presence of three branches of posterior branch of M. This particular pterostigma reminds the pterostigma of the ���hypoperlid��� Idelopsocus, with the important difference in the fact that the pterostigma is covering the area between C and RA but also part of area between RA and RP in Ideloposcus. Hypopsylla differs from the Permopsociidae in the absence of a crossvein between M and CuA. It differs from the Archipsyllidae in the ScP ending in RA instead of going into costal margin (and re-emerging or not distally to reach RA) (Huang et al. 2016 suppl. inf.). The presence of a crossvein between RP and the posterior side of the pterostigma is not discriminant as it is present in the three families (Permopsocus (Permopsocidae), Dichentomum (Psocidiidae) and in the Archipsyllidae). Hypopsylla shares with the Psocidiidae the elongate and narrow areola postica, absence of crossvein between M and CuA, presence of a short crossvein between ScP and costa as in Dichentomum, ScP ending on RA distal of base of RP, but closer to this point than to the pterostigma, unlike to the other taxa of this family. The wing venation of Hypopsylla belmontensis allows its assignment to the Psocidiidae, increasing the diversity of the Permian Permopsocida., Published as part of Prokop, Jakub, Garrouste, Romain & Nel, Andr��, 2016, A new permopsocidan genus and species from the Late Permian of Australia (Insecta: Acercaria: Psocidiidae), pp. 589-592 in Zootaxa 4147 (5) on pages 590-592, DOI: 10.11646/zootaxa.4147.5.7, http://zenodo.org/record/266236, {"references":["Shi, G. R., Waterhouse, J. B. & McLoughlin, S. (2010) The Lopingian of Australasia: a review of biostratigraphy, correlations, palaeogeography and palaeobiogeography. Geological Journal, 45, 230 - 263. http: // dx. doi. org / 10.1002 / gj. 1213","Nel, A., Prokop, J., Nel, P., Grandcolas, P., Huang, Di-ying, Roques, P., Guilbert, E., Dostal, O. & Szwedo, J. (2012) Traits and evolution of wing venation pattern in paraneopteran insects. Journal of Morphology, 273, 480 - 506. http: // dx. doi. org / 10.1002 / jmor. 11036","Tillyard, R. J. (1926) Kansas Permian insects. 8. Copeognatha. American Journal of Science, Series 5, 11, 314 - 349."]}

Genus Eoacridophagus gen. nov. Type species. Eoacridophagus azari sp. nov. (here designated) Diagnosis. Reduced mouthparts with proboscis very short; discal cell open in wing margin; anal area narrow; cell cup closed before wing margin, reaching it by a short stalk; r-m situated well basal of level of apex of R 1, R 2 + 3 not sigmoidal. Etymology. Named after the Eocene period and the genus Acridophagus. Gender masculine., Published as part of Myskowiak, Justine, Garrouste, Romain & Nel, Andre, 2016, A new genus and species of micro bee fly from the Earliest Eocene French amber (Diptera: Mythicomyiidae: Psiloderoidinae), pp. 583-586 in Zootaxa 4114 (5) on page 583, DOI: 10.11646/zootaxa.4114.5.5, http://zenodo.org/record/259075

Genus Eocenotrichia gen. nov. Type species. Eocenotrichia magnifica sp. nov. Etymology. Named after Eocene and ���trichia���, frequent termination of the generic names in the Scenopinidae. Gender feminine. Generic diagnosis. Body length 7.6 mm [female]; head higher than long, flagellum relatively elongate with notched apex for stylus; frons not protruding anteriorly; mouthparts well developed and shorter than head length; R 4 emerging in distal third of cell [r 5]; M 1 joining with R 5, cell [r 5] petiolate to wing margin; costal margin ending at R 5 +M 1; female acanthophorite spines well developed in a marginal row; sternite 8 slightly shorter than tergite 8, posteriorly rounded., Published as part of Garrouste, Romain, Azar, Dany & Nel, Andre, 2016, The oldest accurate record of Scenopinidae in the Lowermost Eocene amber of France (Diptera: Brachycera), pp. 444-450 in Zootaxa 4093 (3) on page 445, DOI: 10.11646/zootaxa.4093.3.10, http://zenodo.org/record/255323

Eocenotrichia magnifica sp. nov. Figs. 1���4 Material. Holotype PA 16841, stored in the Laboratory of Palaeontology, MNHN, Paris, France. Etymology. Named after the excellent preservation of the type specimen. Type horizon. Lowermost Eocene, Sparnacian, level MP 7 of the mammal fauna of Dormaal. Type locality. Le Quesnoy, Chevri��re, region of Creil, Oise department, France. Diagnosis. As for the genus (vide supra); vein R 4 sigmoidal. Description. Body length 7.6 mm [female]. Head 0.75 mm long, 0.96 mm high, higher than long, subspherical, female with broad, raised postocular ridge; antenna elongate, 0.46 mm long, cylindrical, length 0.6 �� head length; antennal style terminal, flagellum 0.3 mm long, notched; frons flat, not protruding anteriorly; mouthparts well developed, 0.42 mm long, distinctly shorter than head length; thorax 1.9 mm long, 1.7 mm high, scutum with dense pile of semi-appressed, very small setae; wing 4.2 mm long, ca. 1.3 mm wide; vein M 1 joining with R 5, cell r 5 petiolate to wing margin, cell r 5 large, 1.8 mm long, 0.4 mm wide; R 5 +M 1 0.2 mm long, ending at wing apex; R 4 sigmoidal, 0.6 mm long, emerging in distal third of cell [r 5]; apex of R 2 + 3 not far from level of base of R 4; vein M 2 absent; vein M 4 originating on discal cell and fused with M 3; costal margin ending at apex of vein R 5 +M 1; cubital veins terminating before wing margin; abdomen elongate and broad, width equal to thorax; abdomen 3.8 mm long, 1.2 mm wide; female genitalia: tergite 10 narrow and band-like, acanthophorite spines present, well developed in a marginal row; sternite 8 slightly shorter than tergite 8, posteriorly rounded. Male unknown. Discussion. Eocenotrichia gen. nov. is placed in the Scenopididae for the wing vein M 4 originating on the discal cell and fused with M 3; in the Scenopidinae for the cubital veins terminating before wing margin, vein M 2 absent, cell [m 1] wide, and in the Metatrichini Winterton & Ware, 2015 for the wing vein M 1 fused to R 5 (Winterton & Ware, 2015). Following the key to scenopinid genera of Winterton & Gharali (2011), within this tribe, Eocenotrichia runs to Propebrevitrichia Kelsey, 1969 in their couplet 25, for the following characters: mouthparts not atrophied; head shorter than high; relatively delicate flies with narrow tapered abdomen; antennal flagellum broad, notched apically; wing with vein R 4 branching from R 5 along distal half of cell [r 5]; female acanthophorite spines present well developed. It shares with Propebrevitrichia, the sister group of all other Metatrichini, the presence of female acanthophorite spines developed in a marginal row (plesiomorphic state for character 28 in Winterton & Ware, 2015), However, Eocenotrichia differs from this genus in that tergite 8 is slightly longer than sternite 8, and the body size is greater than 7 mm instead of being less than 4 mm long (Kelsey, 1969, 1971, 1976; Winterton, 2005). Remarks. As Propebrevitrichia is a South African genus that is the sister group of the clade that comprises all other modern Metatrichini, Winterton & Ware (2015: 23) proposed an African origin of the entire clade during the Late Cretaceous. The present discovery of a Metatrichini in the Earliest Eocene supports the age proposed by these author for this clade. However, the reduced contacts between the African plate and Europe during the period from the Late Cretaceous to the Eocene, questions their hypothesis of an African origin for the Metatrichini. The modern Scenopinidae have predacious larvae associated with wood-boring larvae, termites, woodrat nests, bird���s nests, and carpet beetle larvae (Kelsey, 1981). Birds, termites and Dermestidae are recorded from the Oise amber, suggesting similar biology for Eocenotrichia (Nel & Bourguet, 2006; Kirejtshuk & Nel, 2013)., Published as part of Garrouste, Romain, Azar, Dany & Nel, Andre, 2016, The oldest accurate record of Scenopinidae in the Lowermost Eocene amber of France (Diptera: Brachycera), pp. 444-450 in Zootaxa 4093 (3) on pages 445-446, DOI: 10.11646/zootaxa.4093.3.10, http://zenodo.org/record/255323

Prokop, Jakub, Garrouste, Romain, Nel, André (2016): A new permopsocidan genus and species from the Late Permian of Australia (Insecta: Acercaria: Psocidiidae). Zootaxa 4147 (5): 589-592, DOI: http://doi.org/10.11646/zootaxa.4147.5.7

Hypopsylla gen. nov. Type species. Hypopsylla belmontensis sp. nov. Diagnosis. Wing characters only. ScP ending on RA distal of base of RP, but closer to this point than to the pterostigma (a character also present in Permopsociidae but not in Archipsyllidae); presence of a short crossvein between ScP and costa; a very deep posterior curve of RA in the pterostigma; posterior branch of M with three branches; elongate and narrow areola postica (a character also present in Archipsyllidae but not Permopsociidae), absence of crossvein between M and CuA (a character also present in Archipsyllidae but not Permopsociidae). Etymology. Composite name after Hypoperla and Archypsylla. Gender feminine. Remark. The new genus Hypopsylla (Upper Permian, Australia, H. belmontensis) shares with the Psocidiidae the elongate and narrow areola postica, presence of a short crossvein between ScP and costa as in Dichentomum, ScP ending on RA distal of base of RP, but closer to this point than to the pterostigma, unlike in other taxa of this family., Published as part of Prokop, Jakub, Garrouste, Romain & Nel, Andr��, 2016, A new permopsocidan genus and species from the Late Permian of Australia (Insecta: Acercaria: Psocidiidae), pp. 589-592 in Zootaxa 4147 (5) on page 590, DOI: 10.11646/zootaxa.4147.5.7, http://zenodo.org/record/266236

Family Psocidiidae Tillyard, 1926 Remark. This family currently comprises six valid genera, but only three of them can be accurately considered as Permopsocida, viz. Dichentomum Tillyard, 1926, Stenopsocidium Tillyard, 1935, Hypopsylla gen. nov. Carpenter (1992) synonymized Parapsocidium Zalessky, 1937 with Dichentomum without clear explanation. The other genera of uncertain affinities are Megapsocidium Tillyard, 1935, Austropsocidium Tillyard, 1935, and Liassopsocus Ansorge, 1996. The family is currently known from Australia, possibly Central Europe, Russia, and USA (Tillyard 1926, 1935; Zalessky 1937; Ansorge 1996). Diagnosis (after Huang et al. 2016). Fore- and hindwing with very similar venation; ScP long, ending on RA distal of base of RP in four wings; RP two-branched; M four-branched; areola postica longer than high; no crossvein between M and CuA1. At least Dichentomum has small crossveins between costa and ScP., Published as part of Prokop, Jakub, Garrouste, Romain & Nel, Andr��, 2016, A new permopsocidan genus and species from the Late Permian of Australia (Insecta: Acercaria: Psocidiidae), pp. 589-592 in Zootaxa 4147 (5) on pages 589-590, DOI: 10.11646/zootaxa.4147.5.7, http://zenodo.org/record/266236, {"references":["Tillyard, R. J. (1926) Kansas Permian insects. 8. Copeognatha. American Journal of Science, Series 5, 11, 314 - 349.","Tillyard, R. J. (1935) Upper Permian insects of New South Wales. 3. The order Copeognatha. Proceedings of the Linnean Society of New South Wales, 60, 265 - 279.","Carpenter, F. M. (1992) Treatise on Invertebrate Paleontology, Part R, Arthropoda 4. Geological Society of America Inc, Boulder, 655 pp.","Zalessky, G. M. (1937) Ancestors of some groups of the present day insects. Nature, 140, 847 - 848. http: // dx. doi. org / 10.1038 / 140847 a 0","Ansorge, J. (1996) Insekten aus dem Oberen Lias von Grimmen (Vorpommern, Norddeutschland). Neue Palaontologische Abhandlungen, 2, 1 - 132."]}

Genus Paleoropronia gen. nov. urn:lsid:zoobank.org:act: 214E6B03-C7 DB- 4670-B4D2-427 ABB 5A8A41 Type species Paleoropronia salamonei gen. et sp. nov. Diagnosis See species. Etymology Named after the Paleocene period and Ropronia. Gender feminine., Published as part of Garrouste, Romain, Pouillon, Jean-Marc, Nel, Andr�� & Umr, Isyeb -, 2016, The first Cenozoic roproniid wasp from the Paleocene of Menat, France (Hymenoptera: Proctotrupoidea), pp. 1-9 in European Journal of Taxonomy 239 on page 2, DOI: 10.5852/ejt.2016.239, http://zenodo.org/record/3850241

Paleoropronia salamonei gen. et sp. nov. urn:lsid:zoobank.org:act: D8459184-2211-4B41-B4FE-E72522D78 BCF Figs 1���4 Diagnosis Fore wing venation only. Cell [1M] very narrow, much narrower than cell [2Cu], only 0.2 times as wide as [2Cu]; pterostigma linear, not triangular; base of pterostigma well distal of [1M]; cell [1R1] elongate. Etymology Named after Matt��o Salamone, who helped us to make the photographs and drawing of the type specimen. Type material Holotype FRANCE: MNHN.F. A57266, stored in the collection of Palaeontology (MNHN, Paris). Type stratum and locality FRANCE: Paleocene, spongo-diatomite maar Palaeolake, Menat, Puy-de-D��me. Description A body in rather poor condition with the two fore wings attached. Head 2.3 mm long, 2.8 mm wide, broader than long; head structures hardly visible, except for fragments of the two antennae. Mesosoma 3.9 mm long, 3.6 mm wide; structures hardly visible. Metasoma in poor condition and incomplete but the first segment seems to be narrow tubular, not conical. Fore wing 7.6 mm long, 2.8 mm wide, apparently hyaline except for black pterostigma, 1.7 mm long, 0.23 mm wide, narrow and elongate, situated well distal of cell [1M]; pterostigma and veins covered with small punctuations probably corresponding to the bases of macrotrichiae; cell [1M] quadrangular, 0.7 mm long, 0.1 mm wide, very narrow, much narrower than cell [2Cu]; cell [1R1] very long, 1.9 mm long, 0.4 mm wide; cell [2R1] triangular., Published as part of Garrouste, Romain, Pouillon, Jean-Marc, Nel, Andr�� & Umr, Isyeb -, 2016, The first Cenozoic roproniid wasp from the Paleocene of Menat, France (Hymenoptera: Proctotrupoidea), pp. 1-9 in European Journal of Taxonomy 239 on page 2, DOI: 10.5852/ejt.2016.239, http://zenodo.org/record/3850241

Garrouste, Romain, Pouillon, Jean-Marc, Nel, André, Umr, Isyeb - (2016): The first Cenozoic roproniid wasp from the Paleocene of Menat, France (Hymenoptera: Proctotrupoidea). European Journal of Taxonomy 239: 1-9, DOI: http://dx.doi.org/10.5852/ejt.2016.239

Family Anthracoptilidae Handlirsch, 1922 (sensu Guan et al. 2015 in press) Type genus. Anthracoptilus Lameere, 1917, Late Carboniferous. Other genera. Carrizocladus Rasnitsyn (in Rasnitsyn et al. 2004), Graticladus Novokshonov & Aristov, 2004, Homocladus Carpenter, 1966, Jarmilacladus Rasnitsyn & Aristov, 2004, Mesoptilus Lameere, 1917, Mycteroptila Rasnitsyn, 1977, Paracladus Carpenter, 1966, Spargoptilon Kukalov��, 1965, Strephocladus Scudder, 1885, Strephoneura Martynov, 1940, Strephoptilus Rasnitsyn & Aristov, 2013 (in Aristov et al. 2013), Westphaloptilus Guan et al., 2015. Some genera are currently attributed to the Anthracoptilidae but of uncertain affinities: Adiphlebia Scudder, 1885, Pseudomesoptilus Guan et al., 2015, Rhinomaloptila Rasnitsyn, 1977., Published as part of Nel, Andre, Garrouste, Romain & Prokop, Jakub, 2015, The first African Anthracoptilidae (Insecta: Paoliida) near the Permian ��� Triassic boundary in Kenya, pp. 145-150 in Zootaxa 3925 (1) on page 146, DOI: 10.11646/zootaxa.3925.1.10, http://zenodo.org/record/234742, {"references":["Novokshonov, V. G. & Aristov, D. S. (2004) New taxa of hypoperlids (Insecta: Hypoperlida) from the Upper Permian of the Arkhangelsk Region. Paleontological Journal, 38, 60 - 66.","Carpenter, F. M. (1966) The Lower Permian insects of Kansas. Part 11. The orders Protorthoptera and Orthoptera. Psyche, 73, 46 - 88. http: // dx. doi. org / 10.1155 / 1966 / 47172","Martynov, A. V. (1940) Permian fossil insects from Tshekarda. Trudy Paleontologicheskogo Instituta Akademii nauk SSSR, 11, 1 - 62. [in Russian, with a English summary]","Aristov, D. S., Bashkuev, A. S., Golubev, V. K., Gorochov, A. V., Karasev, E. V., Kopylov, D. S., Ponomarenko, A. G., Rasnitsyn, A. P., Rasnitsyn, D. A., Sinitshenkova, N. D., Sukatsheva, I. D. & Vassilenko, D. V. (2013) Fossil Insects of the Middle and Upper Permian of European Russia. Paleontological Journal, 47, 641 - 832. http: // dx. doi. org / 10.1134 / S 0031030113070010"]}

Afrocladus kenyaensis sp. nov. (Figure 3���4) Material. Holotype specimen NHM II 3069 a, b (print and counterprint), Natural History Museum, London, UK. Etymology. Named after the Kenya country. Age and outcrop. Late Permian (?), Mombasa Basin, Lower unit of Maji ya Chumvi Formation, Duruma Sandstones series, Karoo supergroup, Kenya. Diagnosis. Based on hind (?) wing venation only, wing longer than that of Afrocladus pumilio, RP closely parallel to RA near its base, CuA with four anterior branches instead of three in Afrocladus pumilio. Description. Nearly complete single wing with part of anal area missing, 9.0 mm long, 2.7 mm wide; simple, straight, concave ScP closely parallel to Radius, 0.2 mm apart, ending on costa 3.1 mm from wing base; area between ScP and costal margin narrow, 0.2 mm wide, without visible crossveins or veinlets; radial stem straight, with RP separating from RA 3.2 mm from wing base; RA straight, apparently simple; slightly concave RP with two visible branches; RP closely parallel to RA at its base, touching median vein close to its base; slightly concave median vein divided into two parallel simple branches 2.5 mm from wing base; a brace m-cua between M and CuA 1.8 mm from wing base; CuA and CuP emerging from a common stem; convex CuA anteriorly pectinate with four parallel simple branches less convex than main stem of CuA; concave CuP simple, weakly curved with a rather narrow area between it and CuA, 0.5 mm wide; area between CuP and posterior wing margin ca. 1.1 mm wide; first anal vein convex, simple, weakly curved; second anal vein more poorly preserved but with apparently two posterior branches; two further anal veins more very poorly indicated; several simple crossveins between branches of M and RP; all branches of RP, M and CuA weakly curved and parallel. Discussion. These two fossils fall in the Anthracoptilidae sensu Guan et al. (in press), as it has the following diagnostic characters of the family, i.e. ScP ending on RA or on costal margin; RP emerging from R in basal half of wing; median vein strongly approximate or shortly fused with RP near base of RP (synapomorphy); branches of RP, median vein and CuA strongly parallel at least in their basal halves; median vein braced to CuA by a short convex cross-vein mp-cua; CuA and CuP emerging from a long common stem; main stem of CuA strongly convex, but anteriorly pectinate with at least a series of parallel branches, all less convex than main stem of CuA (synapomorphy); CuP concave simple; at least two anal veins more convex than CuP; anal area can be very large or not. The presence of a common stem Cu from which CuA emerges directly excludes affinities with the archaeorthopteran of the clade Cnemidolestodea Handlirsch, 1937 (sensu B��thoux 2005), in which CuA is also anteriorly pectinate (see discussion in Guan et al. in press). These two wings have a very narrow area between costa and ScP, suggesting they are hindwings, as this area is clearly broader in the forewings of the other Anthracoptilidae. After Rasnitsyn & Aristov (2004), only the genera Jarmilacladus, Rhinomaloptila, and Spargoptilon share with Afrocladus the ScP ending on costal margin and not on radius. This character seems to be rather variable at generic level because Mesoptilus carpenteri Guan et al., 2015, shares it while Mesoptilus dolloi (Lameere 1917) has ScP ending on radius. Afrocladus differs from all anthracoptilids except Paracladus retardatus Carpenter, 1966, in the distinctly shorter ScP, with apex close to base of RP. Afrocladus pumilio and Afrocladus kenyaensis have also very small wings, 5.5 to 9.0 mm long, unlike the other Anthracoptilidae that have wings ranging between 17 to 48 mm long. Conculusion The presence of fossil insects on the surface of borehole samples is highly improbable, although few other records are known (viz. the case of the holotype of Permophlebia uralica Nel et al., 2001). These two discoveries in the same outcrop suggest that the corresponding layers are probably very rich in fossil terrestrial arthropods. It would be of great interest to search for new material in situ. Afrocladus pumilio and Afrocladus kenyaensis are the first African and the youngest representatives of the Anthracoptilidae, and of the Paoliida. They are of very small sizes, distinctly smaller than all the other Paoliida except some Blattinopsidae (Glaphyrophlebia pygmaea (Meunier, 1907) from the Late Carboniferous of France, with a wing 14 mm long) (H��rnschemeyer & Stapf 2001). This size reduction could be put in parallel to a similar phenomenon that seems to have affected the last representatives of the order Palaeodictyoptera during the Late Permian (B��thoux et al. 2007). It could be due to adaptations to environmental changes that have taken place around the P-T boundary (Aristov et al. 2013)., Published as part of Nel, Andre, Garrouste, Romain & Prokop, Jakub, 2015, The first African Anthracoptilidae (Insecta: Paoliida) near the Permian ��� Triassic boundary in Kenya, pp. 145-150 in Zootaxa 3925 (1) on pages 147-148, DOI: 10.11646/zootaxa.3925.1.10, http://zenodo.org/record/234742, {"references":["Bethoux, O. (2005) Cnemidolestodea (Insecta): an ancient order reinstated. Journal of Systematic Palaeontology, 3, 403 - 408. http: // dx. doi. org / 10.1017 / S 147720190500163 X","Carpenter, F. M. (1966) The Lower Permian insects of Kansas. Part 11. The orders Protorthoptera and Orthoptera. Psyche, 73, 46 - 88. http: // dx. doi. org / 10.1155 / 1966 / 47172","Nel, A., Bethoux, O., Bechly, G., Martinez-Delclos, X. & Papier, F. (2001) The Permo-Triassic Odonatoptera of the ' protodonate' grade (Insecta: Odonatoptera). Annales de la Societe Entomologique de France, New Series, 37, 501 - 525.","Hornschemeyer, T. & Stapf, H. (2001) Review of Blattinopsidae (Prothortoptera) with description of new species from the Lower Permian of Niedermoschel (Germany). Neue Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 221, 81 - 109.","Bethoux, O., Nel, A., Schneider, J. W. & Gand, G. (2007) Lodetiella magnifica nov. gen. and nov. sp. (Insecta: Palaeodictyoptera; Permian), an extreme situation in wing morphology of palaeopterous insects. Geobios, 40, 181 - 189. http: // dx. doi. org / 10.1016 / j. geobios. 2006.04.001","Aristov, D. S., Bashkuev, A. S., Golubev, V. K., Gorochov, A. V., Karasev, E. V., Kopylov, D. S., Ponomarenko, A. G., Rasnitsyn, A. P., Rasnitsyn, D. A., Sinitshenkova, N. D., Sukatsheva, I. D. & Vassilenko, D. V. (2013) Fossil Insects of the Middle and Upper Permian of European Russia. Paleontological Journal, 47, 641 - 832. http: // dx. doi. org / 10.1134 / S 0031030113070010"]}

Afrocladus pumilio sp. nov. (Figure 1���2) Material. Holotype specimen NHM II 3070, Natural History Museum, London, UK. Etymology. Named after the very small size of this wing. Age and outcrop. Late Permian (?), Mombasa Basin, Lower unit of Maji ya Chumvi Formation, Duruma Sandstones series, Karoo supergroup, Kenya. Diagnosis. Based on hind (?) wing venation only, wing shorter than that of Afrocladus kenyaensis, RP not closely parallel to RA near its base, CuA with only three anterior branches instead of four in Afrocladus kenyaensis. Description. Single wing with costo-apical part missing, 5.5 mm long, 1.76 mm wide; simple, straight, concave ScP closely parallel to Radius, 0.1 mm apart, ending on costa 2.6 mm from wing base; area between ScP and costal margin narrow, 0.2 mm wide, without visible crossveins or veinlets; radial stem straight, with RP separating from RA 2.1 mm from wing base; RA straight, with apex missing; slightly concave RP with two visible branches; RP touching median vein near its base; slightly concave median vein divided into two parallel branches 1.7 mm from wing base; a brace m-cua between M and CuA 1.1 mm from wing base; CuA and CuP emerging from a common stem; convex CuA anteriorly pectinate with three parallel simple branches less convex than main stem of CuA; concave CuP simple, weakly curved with a rather narrow area between it and CuA, 0.35 mm wide; area between CuP and posterior wing margin 0.7 mm wide; first anal vein convex, simple, weakly curved; second anal vein with two posterior branches; two further anal veins more weakly indicated; several simple crossveins present between branches of M and RP; all branches of RP, M and CuA weakly curved and parallel., Published as part of Nel, Andre, Garrouste, Romain & Prokop, Jakub, 2015, The first African Anthracoptilidae (Insecta: Paoliida) near the Permian ��� Triassic boundary in Kenya, pp. 145-150 in Zootaxa 3925 (1) on pages 146-147, DOI: 10.11646/zootaxa.3925.1.10, http://zenodo.org/record/234742

Garrouste, Romain, Nel, André (2015): New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata). Zootaxa 4028 (3): 354-366, DOI: http://dx.doi.org/10.11646/zootaxa.4028.3.2

Genus Afrococladus gen. nov. Type species. Afrocladus pumilio sp. nov., other species. Afrocladus kenyaensis sp. nov. Diagnosis. Hind (?) wing characters only. Wings very small (less than 9 mm long, instead of ranging between 17 to 48 mm in other Anthracoptilidae), with ScP very short, ending on costal margin at or close the level of base of RP.

Nel, Andre, Garrouste, Romain, Prokop, Jakub (2015): The first African Anthracoptilidae (Insecta: Paoliida) near the Permian—Triassic boundary in Kenya. Zootaxa 3925 (1): 145-150, DOI: http://dx.doi.org/10.11646/zootaxa.3925.1.10

Petrolestes hendersoni Cockerell, 1927 (Figs. 1–5) Material. holotype specimen UCM 15619, University of Colorado, Boulder, Geology Section, Museum Annex, USA; new specimen USNM without number, Smithsonian Institution, National Museum of Natural History, USA. Age and outcrop. Eocene, Green River Formation; the type specimen UCM 15619 comes from the Ute Trail, Roan Plateau, Garfield County; the new specimen USNM without number comes from Wighton Garfield County; both in Colorado, USA. Emended diagnosis. Wing characters only. Intercalaries between MA and MP present just distal of nodus level; 17 postnodals; wing rather large, 8.0 mm wide at nodus level. Description of new specimen. Body with head and five basal segment of abdomen attached to thorax, together with the four wings; only the left forewing is nearly complete, basal half of left hind wing and basal fourth of right wings preserved; no trace of secondary genital structures on abdominal segment 2 (female specimen). Head rather broad, 3.6 mm long, 5.2 mm wide, eyes 3.0 mm wide, widely separated, 2.4 mm apart; ocelli not discernable; thorax 6.0 mm high, moderately prognathous, thoracic skewness (sensu Needham and Anthony, 1903) moderate, comparable to that of a modern Aeshnidae, angle x 35 °, angle z ca. 30 °; interpleural suture (sensu Asahina 1957) incomplete but formed at dorsal 1 / 2; abdomen 2.0 mm wide. Legs robust, with tibia spines strong, slightly longer than distance between two spines. Wings hyaline, forewing ca. 40.0 mm long; max. width 8.5 mm, wing 8.0 mm wide at nodus level; a very short petiole, 1.5 mm long; anal area with one row of cells between AA and AP; subdiscoidal cell broad elongate, 1.8 mm long, 0.8 mm wide; distance from base to arculus 5.0 mm, from arculus to nodus 11.2 mm, from nodus to pterostigma 16.5 mm; nodus in a basal position; a significant increase of spine density at the apical costal margin; pterostigma long and broad, 4.0 mm long, 0.8 mm wide, covering six cells; pterostigmal brace perpendicular to RA and RP 1; distance from wing base to Ax 1 3.8 mm, from Ax 1 to Ax 2 1.5 mm, Ax 2 nearly aligned with arculus; no supplementary antenodal crossvein; discoidal cell unicellular, 1.4 mm long, 1.4 mm wide, distinctly broadened in its distal part, basal side 0.8 mm, anterior side 1.2 mm, posterior side 1.4 mm, distal side MAb 1.4 mm; ScP not crossing through nodus as in Stenolestes Scudder, 1895; nodal crossvein very oblique; subnodus slightly oblique with corresponding crossvein below (between RP 1 / 2 and IR 2) of inverted obliquity; ca. 17 postnodal crossveins preserved, not aligned with the 16 postsubnodal crossveins; bases of RP 3 / 4 and IR 2 between arculus and nodus, closer to nodus than to arculus, base of RP 3 / 4 6.5 mm from arculus and 4.7 mm from nodus; base of RP 2 two cells, 2.2 mm distal of subnodus; base of IR 1 four cells and 4.0 mm distally; oblique crossvein ‘O’ absent; cubito-anal area with five rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin well distal of nodus level; CuA weakly curved; postdiscoidal area with one row of cells up to nodus level and two rows more distally; area between RP 3 / 4 and MA with at least two secondary longitudinal veins in between; three rows of cells between RP 3 / 4 and IR 2 and one row between RP 1 and IR 1; three rows of cells between IR 1 and RP 2 opposite pterostigma, but only one basally. Preserved part of hind wing nearly identical to forewing, even the shape of the discoidal cell nearly the same. Only the position of the nodus is slightly different, as distance from arculus to nodus is 8.7 mm. Discussion. The comparison of the preserved parts (hind wing basal two-thirds) of the venation of the type of Petrolestes hendersoni with that of this new fossil shows that they are nearly identical in every point (petiole length, nodus, subnodus, discoidal cell, antenodal and cubito-anal areas, relative positions of bases of branches of RP, number of rows of cells between main veins in basal part of wings). Also the dimensions of the wing structures are nearly identical in the two specimens. As both specimens come from the Green River Formation in Colorado and even from outcrops in the same county, we consider that they belong to the same species. The new specimen allows more precision in several points concerning this taxon, viz. it has: no oblique vein ‘O’, a long pterostigma covering numerous cells, a long zigzagged IR 1, a short petiole, and the discoidal cells of the fore and hind wings are nearly identical. These characters confirm the position of Petrolestes close to Congqingia Zhang, 1992 in the Dysagrioninae, rather than close to the Sieblosiidae. In particular Petrolestes has not the subnodal veinlet vertical or anteriorly oriented and its nodus is not traversed by ScP, unlike in the Sieblosiidae; it also has no oblique vein ‘O’. Petrolestes has also a significant increase of spine density at the apical costal margin, a character proposed by Bechly (1996) as a potential synapomorphy of the Zygoptera, absent in the Sieblosiidae (Fleck et al. 2004). Petrolestes and Congqingia share with Dysagrion a rather unique shape of the discoidal cell, potential synapomorphy even if it is also present in the Sieblosiidae. The presence of a ‘basal costal margin between wing base and nodus distinctly convex curved’ (a character absent in the Sieblosiidae, see Nel et al. 2005), plus the ‘antesubnodal space without any crossveins’ (a character present in many Zygoptera and in the Sieblosiidae) would be the unique putative synapomorphies of the Dysagrioninae with the modern Thaumatoneuridae, after Bechly (2014). If Dysagrion shares with Thaumatoneura similar relative positions of the nodus and bases of the branches of RP, it is not the case for Petrolestes and Congqingia, which have the bases of RP 3 / 4 and IR 2 basally recessed midway between the nodus and the arculus. Also the discoidal cell of Thaumatoneura is elongate and much narrower than in all the taxa currently included in the Dysagrioninae. Also Thaumatoneura has long spines on its legs, unlike Petrolestes, and its thoracic skewness is much more pronounced. Thus the attribution of this group of damselflies as sister group of the modern Thaumatoneuridae remains debatable. Several characters suggest that Petrolestes (and consequently Congqingia and maybe Dysagrion) could not be a Zygoptera: the head appears not as transverse as in a Zygoptera (but this could be due to the compression of the fossil that can strongly deform the head shape; similar situations occur for fossil Lestes in compression on rocks from the Oligocene of France, so that the head seems to be less transverse than it was certainly in the living animal); the thorax is not very skewed, similarly to what happen in Epiproctophora (but this could be due to the robustness of this animal); the legs have spines shorter and stronger than in Zygoptera; this character is rather present in Epiproctophora. Also the position of the arculus near Ax 2 instead of being midway between Ax 2 and Ax 1 or closer to Ax 1 is a plesiomorphic character that would exclude a position within the Epiproctophora for Petrolestes. The presence of an incomplete interpleural suture but rather long and formed at dorsal ½ is a plesiomorphic character present in Zygoptera (the interpleural suture is complete in Thaumatoneura) but not in Epiproctophora. It remains that the thoracic skewness and relatively short leg spines of Petrolestes suggest affinities with the Epiproctophora rather than with Zygoptera. Petrolestes, and consequently Congqingia and Dysagrion, could be taxa in the stem lineage of the Epiproctophora rather than Zygoptera. It would be more accurate to have information on the genital appendages of these damselflies before an attribution close to the Epiproctophora. Petrolestes differs from Congqingia in the longer pterostigma, covering six cells, instead of two, in the broader area between IR 1 and RP 2 with longer intercalary veins between them, and in the broader cubito-anal area with five-six rows of cells instead of two (Zhang 1992)., Published as part of Garrouste, Romain & Nel, André, 2015, New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata), pp. 354-366 in Zootaxa 4028 (3) on pages 355-359, DOI: 10.11646/zootaxa.4028.3.2, http://zenodo.org/record/232682, {"references":["Needham, J. G. & Anthony, M. H. (1903) The skewness of the thorax in the Odonata. Journal of the New York Entomological Society, 11, 117 - 123.","Asahina, S. (1957) On the pterothoracic interpleural suture of Mesozoic and Recent Odonata. J ournal of the Faculty of Sciences, Hokkaido University, Zoology, (6), 13, 1 - 7.","Bechly, G. (1996) Morphologische Untersuchungen am Flugelgeader der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Berucksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Special Volume 2, 402 pp.","Fleck, G., Bechly, G., Martinez-Delclos, X., Jarzembowski, E. A. & Nel, A. (2004) A revision of the Mesozoic dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Odonatoptera: Panodonata). Geodiversitas, 26, 33 - 60.","Nel, A., Petrulevicius, J. F., Gentilini, G. & Martinez-Delclos, X. (2005) Phylogenetic analysis of the Cenozoic family Sieblosiidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. http: // dx. doi. org / 10.1016 / j. geobios. 2003.10.007","Bechly, G. (2014) Phylogenetic systematics of Odonata. Available from: http: // members. tripod. de / GBechly / phylosys. htm (Accessed 7 Oct. 2015)"]}

Pseudostenolestes bechlyi sp. nov. (Figs. 8–13) Material. Holotype specimen MeI 14781 a–b, paratypes specimens MeI 13209, MeI 14712, and MeI 14794 a–b, all at SMF. Etymology. Named after our friend and colleague Günter Bechly, for his contributions in odonatology. Age and outcrop. Lower Middle Eocene, Geiseltalian (Lower Lutetian), ca. 47 million year old former Maar lake Messel, near Darmstadt, Hesse, Germany. Diagnosis. As for the family, plus presence of a dark metallic zone crossing the wing around nodus level. Descriptions. Specimen MeI 14781 a–b: MeI 14781 a (a hind wing base up to nodus level): length of fragment 26.8 mm, max. width 8.9 mm; MeI 14781 b (apical two-thirds of wing): 25.7 mm long; max. width 8.2 mm; wing hyaline except for a broad dark area crossing it around nodus; wing ca. 32.3 mm long, 8.9 mm wide at nodus level; a very short petiole, 0.7 mm long; anal area with two rows of cells between AA and AP because of the presence of a strong vein emerging from posterior side of subdiscoidal cell in its middle and directed towards wing base; subdiscoidal cell broad and elongate, 1.2 mm long, 0.6 mm wide; CuP vein obliquely oriented between AA and MP+CuA; distance from base to arculus 2.9 mm, from arculus to nodus 8.6 mm, from nodus to pterostigma 12.3 mm; nodus in a basal position; pterostigma long and broad, 5.2 mm long, 0.8 mm wide, covering ca. four cells; pterostigmal brace absent; distance from wing base to Ax 1 2.8 mm, from Ax 1 to Ax 2 1.8 mm, Ax 1 aligned with arculus; at least three supplementary antenodal crossveins of first row between C and ScP distal of Ax 2, without corresponding crossvein of second row; discoidal cell unicellular, 1.6 mm long, 1.2 mm wide, distinctly broadened in its distal part, basal side 0.6 mm long, anterior side 1.2 mm long, posterior side 1.1 mm long, distal side MAb 1.2 mm long; ScP not crossing through nodus, unlike in Stenolestes Scudder, 1895; nodal crossvein oblique; subnodus slightly oblique with corresponding crossvein below (between RP 1 / 2 and IR 2) of inverted obliquity; ca. 17 postnodal crossveins preserved, not aligned with the 18 postsubnodal crossveins; bases of RP 3 / 4 and IR 2 between arculus and nodus, closer to arculus than to nodus; base of RP 3 / 4 3.6 mm from arculus; base of RP 2 aligned with subnodus; base of IR 1 five cells and 3.3 mm distally; oblique crossvein ‘O’ present two cells distal of base of RP 2; cubito-anal area with four-six rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin just distal of nodus level; CuA strongly curved; area between MP and posterior wing margin very broad; postdiscoidal area distally widened, with one or two secondary longitudinal veins between MP and MA; area between RP 3 / 4 and MA with two secondary longitudinal veins in between; four rows of cells between RP 3 / 4 and IR 2 and two rows between RP 1 and IR 1; three rows of cells between IR 1 and RP 2 opposite pterostigma, but only one basally. Specimen MeI 13209 (a nearly complete hind wing). Hyaline except for a metallic colored area crossing it around nodus; 34.3 mm long, 9.4 mm wide at nodus level; length of metallic colored area at costal margin 6.9 mm; nearly all preserved details of venation identical to those of specimen MeI 14781 a–b. Specimen MeI 14712 (distal two-thirds of a wing). Hyaline except for a metallic colored area around nodus. Preserved part 24.0 mm long; length of metallic colored area at costal margin 6.9 mm; all preserved parts are identical to those of specimen MeI 14781 a–b. Specimen MeI 14794 a–b: fragment MeI 14794 a (nearly complete wing) 34.3 mm long, 9.3 mm wide; MeI 14794 b (fragment of median part of wing): max. width 9.2 mm. All preserved parts and pattern of coloration are identical to those of specimen MeI 14781 a–b, except for the distinctly less oblique subnodus. Discussion. The comparable preserved parts of these wings are nearly identical, supporting their attribution to the same taxon. The presence of discoidal cells of ‘sieblosiid type’ together with the positions of the bases of RP 3 / 4 and IR 2 midway between arculus and nodus suggest possible affinities with the Sieblosiidae or with the Dysagrioninae: Petrolestini. The presence of the oblique vein ‘O’ together supports affinities with the Sieblosiidae rather than with the Dysagrioninae. Nevertheless, Pseudostenolestes strongly differs from the known Sieblosiidae in the following characters: shape of nodus with ScP not crossing through it; base of RP 2 only one or two cells distal of subnodus instead of being more than five cells distal of it; presence of secondary antenodal crossveins distal of Ax 2. Furthermore Pseudostenolestes differs from both the Sieblosiidae and the Dysagrioninae (but also from all Zygoptera) in the very particular organization of the cubito-anal veins, viz. there is a strong vein emerging from posterior side of subdiscoidal cell in its middle and directed towards wing base. This structure constitutes one of the main synapomorphies of the Epiproctophora (Fleck et al., 2003; Bechly, 2014). It corresponds to the vein CuAb that is separating from CuAa in the postero-distal angle of the subdiscoidal cell, and is distally meeting with AA to vanish in the basal part of cubito-anal area. This structure is clearly visible in the Heterophlebioptera Bechly, 1996, in the Epiophlebiidae, (less distinct in the Stenophlebiidae), the Isophlebioptera, but also in the anisopteran Liassogomphidae (see Fleck et al. 2003). Fleck et al. (2004) proposed that the Sieblosiidae should be removed from the Zygoptera and could be the sister group of the Epiproctophora. Pseudostenolestes shares with the Epiproctophora the synapomorphic presence of a strong CuAb directed towards wing base. Pseudostenolestes has a subdiscoidal cell broader than in the Zygoptera and the Sieblosiidae. This character is reminiscent of the situation in the Isophlebioptera in which the hind wing subdiscoidal cell is enlarged and has a bulged posterior margin (Nel et al., 1993; Bechly, 1996, 2014). Nevertheless, nearly all the representatives of this clade have subdiscoidal cells broader than in Pseudostenolestes and a different pattern of the longitudinal veins. Only the Cyclothemistidae, Bechly, 1996 (e.g., Cyclothemistinae Cyclothemis Pritykina, 1980, and Shurabiola Pritykina, 1980, and Pseudotriassothemistinae Pseudotriassothemis), have a subdiscoidal cell as narrow as Pseudostenolestes. Interestingly the Cyclothemistidae also share with Pseudostenolestes several other characters, viz. the distal side (MAb) of the discoidal cell is about twice as long as the basal side (= posterior part of arculus); the typical pattern of the longitudinal veins in Isophlebioptera is reversed (the same situation also occurs in the Cyclothemistidae Handlirsch, 1939); thus the postdiscoidal space is not narrowed and RP 3 / 4 is not parallel to IR 2. If the Cyclothemistinae have a narrower cubito-anal area than that of Pseudostenolestes, the Triassic Pseudotriassothemis shares with Pseudostenolestes a broad cubito-anal area. Furthermore the CuP vein of Pseudostenolestes is obliquely oriented between AA and MP+CuA. This tendency is achieved in Pseudotriassothemis, in which CuP appears aligned with basal part of AA (Fujiyama 1991). Pseudostenolestes differs from the Cyclothemistidae in the subnodus aligned with the nodal crossing, instead of being in a more distal position (Pritykina 1980; Fujiyama 1991). This important difference is sufficient to justify a family separation. Mesozoic representatives of the Isophlebioptera have the discoidal cells of the fore and hind wings of different shapes (see Nel et al. 1993). Thus we can suppose that the specimens MeI 14781 a–b and MeI 13209 correspond to hind wings with closed discoidal cells. Pseudostenolestes can be considered as the first Cenozoic and youngest Isophlebioptera, while this clade was previously known only between the Upper Triassic and the Lower Cretaceous. This implies a position of ‘ghost’ taxa of the Pseudostenolestidae during the Upper Cretaceous. Until now, if we except the Cenozoic Sieblosiidae, the Epiophlebioptera was the unique Mesozoic subclade (except the Anisoptera) of the Epiproctophora that survived the Cretaceous-Cenozoic mass extinction, as it is known by modern species and a very recently discovered taxon in the mid Cretaceous Burmese amber (Bechly and Poinar 2013). The present discovery of an Isophlebioptera in the Paleogene demonstrates that this clade survived the major odonatological turnover of the Cenomanian-Turonian (Nel et al. 2010) and the K-T extinction., Published as part of Garrouste, Romain & Nel, André, 2015, New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata), pp. 354-366 in Zootaxa 4028 (3) on pages 361-364, DOI: 10.11646/zootaxa.4028.3.2, http://zenodo.org/record/232682, {"references":["Fleck, G., Bechly, G., Martinez-Delclos, X., Jarzembowski, E. A., Coram, R. & Nel, A. (2003) Phylogeny and classification of the Stenophlebioptera (Odonata, Epiproctophora). Annales de la Societe Entomologique de France, (N. S.), 39, 55 - 93.","Bechly, G. (2014) Phylogenetic systematics of Odonata. Available from: http: // members. tripod. de / GBechly / phylosys. htm (Accessed 7 Oct. 2015)","Bechly, G. (1996) Morphologische Untersuchungen am Flugelgeader der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Berucksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Special Volume 2, 402 pp.","Fleck, G., Bechly, G., Martinez-Delclos, X., Jarzembowski, E. A. & Nel, A. (2004) A revision of the Mesozoic dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Odonatoptera: Panodonata). Geodiversitas, 26, 33 - 60.","Pritykina, L. N. (1980) New fossil Odonata from the Mesozoic of Central Asia. In: Dolin, V. G., Panfilov, D. V., Ponomarenko, A. G., and Pritykina, L. N. (Eds.), Mesozoic fossil insects. Kiev: Naukova Dumka, pp. 1 - 134. [in Russian]","Fujiyama, I. (1991) Late Triassic insects from Mine, Yamaguchi, Japan. Part 1. Odonata. Bulletin of the National Science Museum, (C), 17, 9 - 56.","Bechly, G. & Poinar, G. O. Jr. (2013) Burmaphlebia reifi gen. et sp. nov., the first anisozygopteran damsel-dragonfly (Odonata: Epiophlebioptera: Burmaphlebiidae fam. nov.) from Early Cretaceous Burmese amber. Historical Biology, an International Journal of Paleobiology, 25, 233 - 237. http: // dx. doi. org / 10.1080 / 08912963.2012.753884","Nel, A., Nel, P., Petrulevicius, J. F., Perrichot, V., Prokop, J. & Azar, D. (2010) The Wagner Parsimony using Morphological Characters: a new method for palaeosynecological studies. Annales de la Societe Entomologique de France, (N. S.), 46, 276 - 292."]}

Petrolestes messelensis sp. nov. (Figs. 6–7) Material. Holotype specimen MeI 11777 (the distal two-third of a wing), SMF. Age and outcrop. Lower Middle Eocene, Geiseltalian (Lower Lutetian), ca. 47 million year old former Maar lake Messel, near Darmstadt, Hesse, Germany. Diagnosis. Wing characters only. Intercalaries between MA and MP very short and only present close to posterior wing margin; 15 postnodals; wing rather small, 5.4 mm wide at nodus level. Etymology. Named after the Messel Formation. Description. Wing hyaline in its preserved part; fragment 17.8 mm long, 5.4 mm wide at nodus; distance from nodus to pterostigma ca. 10.4 mm; pterostigma long and broad, 1.8 mm long, 0.8 mm wide, covering ca. four-six cells; pterostigmal brace not oblique; no supplementary antenodal crossvein of first row or second row in preserved part of antenodal space; ScP not crossing through nodus as in Stenolestes Scudder, 1895; nodal crossvein oblique; subnodus nearly vertical with crossvein between RP 1 / 2 and IR 2 below it of inverted obliquity; 15 postnodal crossveins preserved, not aligned with the 14 postsubnodal crossveins; bases of RP 3 / 4 and IR 2 between arculus and nodus, probably closer to arculus than to nodus (RP 3 / 4 and IR 2 are basally ‘mixed’ due to taphonomic deformation); base of RP 2 three cells distal of subnodus; base of IR 1 four cells distally; oblique crossvein ‘O’ absent; cubito-anal area with four-five rows of cells between CuA and posterior wing margin; CuA reaching posterior wing margin well distal of nodus level; CuA weakly curved; area between MP and posterior wing margin moderately broad; postdiscoidal area not widened distally, with very short secondary longitudinal vein between MP and MA near posterior wing margin; area between RP 3 / 4 and MA with three secondary longitudinal veins in between; four rows of cells between RP 3 / 4 and IR 2, two rows between RP 1 and IR 1; three rows of cells between IR 1 and RP 2 opposite pterostigma, but only one basally. Discussion. Although incompletely preserved, this wing is very similar to that of the new specimen of Petrolestes hendersoni, especially in the long pterostigma, with brace not oblique, bases of RP 3 / 4 and IR 2 well basal of subnodus, absence of oblique vein ‘O’, organisation of the intercalaries between main veins, position of the bases of RP 3 / 4 and IR 2, shape of the subnodus and of the crossveins situated below it. Thus this new fossil can be accurately attributed to the genus Petrolestes. It differs from P. hendersoni in few characters, viz. the intercalaries between MA and MP are very short only present close to the posterior wing margin instead of being present just distal of nodus level; and 15 postnodals instead of 17; wing distinctly smaller as its width at nodus is 5.4 mm instead of 8.0– 8.1 mm for P. hendersoni., Published as part of Garrouste, Romain & Nel, André, 2015, New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata), pp. 354-366 in Zootaxa 4028 (3) on page 359, DOI: 10.11646/zootaxa.4028.3.2, http://zenodo.org/record/232682

Genus Pseudostenolestes gen. nov. Type species. Pseudostenolestes bechlyi sp. nov. Diagnosis. As for the family. Etymology. Named after Stenolestes, type genus of the Sieblosiidae to indicate the great similarities with the Sieblosiidae. Gender masculine., Published as part of Garrouste, Romain & Nel, André, 2015, New Eocene damselflies and first Cenozoic damsel-dragonfly of the isophlebiopteran lineage (Insecta: Odonata), pp. 354-366 in Zootaxa 4028 (3) on page 360, DOI: 10.11646/zootaxa.4028.3.2, http://zenodo.org/record/232682

Urolibellula eocenica sp. n. (Figs. 1���3) Material. Holotype specimen without number, labelled ���Dragonfly, Anisoptera gen. Corduliidae ���, USNM (United State National Museum). Age and outcrop. Eocene, Green River Formation, Parachute Creek Member, 43���47 Myrs, Wighton, Garfiel County, Colorado. Diagnosis. As for the genus. Description. Two forewings, one hind wing, and three legs attached to fragments of the thorax. Wings hyaline, pterostigmata dark brown. Forewing length 38.0 mm; width at nodus 8.0 mm; distance from base to nodus 20.0 mm; nodus situated distinctly closer to apex than to base; distance from nodus to pterostigma 11.0 mm; distance from base to arculus 6.0 mm; primary antenodal crossveins Axl and Ax 2 aligned but not stronger than other 10 secondary antenodals (all complete except most distal one, but not well aligned with antenodals of second row between ScP and RA); Axl 2.0 mm basal of arculus and Ax 2 5.0 mm distal of Axl, a long ���cordulegastrid gap��� and a gap directly distal of arculus; seven postnodal crossveins between nodus and pterostigma not aligned with six postsubnodal crossveins; a ���libellulid gap��� directly distal of subnodus concerning one postnodal; pterostigma 4.0 mm long and 1.0 mm wide; pterostigma with a basal brace not aligned with its basal side, distally displaced and weakly oblique; pterostigma covering two cells; arculus nearly totally straight and aligned with basal side of discoidal triangle; origins of RP and MA (sectors of arculus) separated at their extreme bases in arculus; free portion of RP +MA longer than posterior crossvein in arculus; hypertriangle 3.8 mm long and 1.0 mm wide, free; its costal side (MA) weakly curved; discoidal triangle not transverse but broad, divided into three cells by crossveins touching in one point; length of basal side of discoidal triangle 3 mm, length of costal side 2.8 mm, length of distal side MAb 4 mm; MAb straight; pseudo-anal vein PsA distinct; subdiscoidal triangle three-celled; basal space free; submedian space only crossed by CuP; anal area max. 2.0 mm wide with two rows of cells; cubito-anal area max. 4.5 mm wide with up to five rows of cells; MP weakly curved and ending just distal of nodus; area between CuA and MP not widened; basal part of postdiscoidal area with 2���3 rows of cells; postdiscoidal area not widened distally (width near discoidal triangle 2.5 mm; width at hind margin 2.3 mm); Mspl nearly straight, ending at MAa, with one row of cells between it and MAa; MAa weakly curved; RP 3 / 4 and MA weakly undulate and closely parallel with only one row of cells between them up to hind margin; first branching of RP 6.1 mm basal of subnodus; IR 2 originating on RPl/ 2; RP 2 aligned with subnodus; only one lestine oblique vein ��� O ��� between RP 2 and IR 2, 0.5 mm and one cell distal of subnodus; presence of a post-oblique-vein gap; second crossvein between RP 1 and RP 2 developed as a ���libellulid oblique vein;��� only one bridge crossvein between RP 2 and IR 2 basal of subnodus; area between RP 2 and IR 2 distally widened, but only one row of cells between these veins, except near hind margin (two rows of cells); a well-defined Rspl with one row of cells between it and IR 2; RP 2 weakly undulate; RPl and RP 2 basally parallel with only one row of cells between them, but about 1.0 mm basal of pterostigma, area between these veins widening progressively; a well-defined pseudo-IRl originating on RPl beneath pterostigma; three rows of cells between pseudo-IRl and RPl and between pseudo-IRl and RP 2; no apical supplement (strong concave supplementary sector developed between RP 1 and RP 2). Hindwing: length 37.0 mm; width at nodus 15.0 mm; distance from base to nodus 15.0 mm; nodus situated distinctly closer to base than to apex; distance from nodus to pterostigma 15.5 mm; distance from base to arculus 5.5 mm; primary antenodal crossveins Axl and Ax 2 aligned but not stronger than other five antenodals (all complete); Axl 3 mm basal of arculus and Ax 2 is 5.5 mm distal of Axl; only four antesubnodal crossveins, with a long ���cordulegastrid gap��� and a gap directly distal of arculus; 10 postnodal crossveins between nodus and pterostigma not aligned with eight postsubnodal crossveins; a ���libellulid gap��� directly distal of subnodus concerning two postnodals; pterostigma 3.5 mm long and 1.0 mm wide; pterostigma with a basal brace not aligned with its basal side, distally displaced and weakly oblique; pterostigma covering two cells; arculus nearly totally straight and aligned with basal side of discoidal triangle; origins of RP and MA (sectors of arculus) separated at their extreme bases in arculus; free portion of RP +MA longer than posterior crossvein in arculus; hypertriangle 3.0 mm long and 0.75 mm wide, free; its costal side (MA) distinctly curved; discoidal triangle not transverse but broad, and free; length of basal side of discoidal triangle 1.8 mm, length of costal side 3.0 mm, length of distal side MAb 3.0 mm; MAb straight; pseudo-anal vein PsA distinct; subdiscoidal triangle unicellular; basal space free; submedian space only crossed by CuP; anal area max. 12.5 mm wide with about ten rows of cells; cubito-anal area max. 8.0 mm wide with up to six rows of cells; CuA strongly sigmoidally curved and with a strong CuAb; anal loop long and narrow, posteriorly closed and with a concave well-defined midrib with a distinct toe; MP strongly curved and ending just basal of nodus; area between CuA and MP not widened; basal part of postdiscoidal area with only two rows of cells; postdiscoidal area strongly widened distally (width near discoidal triangle 3.5 mm; width at hind margin 7.0 mm); a Mspl nearly straight ending on MAa, with one row of cells between it and MAa; MAa weakly curved but with a strong bend near posterior wing margin; RP 3 / 4 and MA not undulate and closely parallel with only one row of cells between them up to hind margin; first branching of RP 5.0 mm basal of subnodus; IR 2 originating on RPl/ 2; RP 2 aligned with subnodus; only one lestine oblique vein ��� O ��� between RP 2 and IR 2, 0.5 mm and one cell distal of subnodus; presence of a post-oblique-vein gap; second crossvein between RP 1 and RP 2 developed as a ���libellulid oblique vein;��� only one bridge crossvein between RP 2 and IR 2 basal of subnodus; area between RP 2 and IR 2 distally widened, but only one row of cells between these veins, except near hind margin (two rows of cells); a well-defined Rspl with one row of cells between it and IR 2; RP 2 weakly undulate; RPl and RP 2 basally parallel with only one row of cells between them, but about 1.5 mm basal of pterostigma, area between these veins widening progressively; a well-defined pseudo-IRl originating on RPl beneath pterostigma; three rows of cells between pseudo-IRl and RPl and between pseudo-IRl and RP 2; no apical supplement (strong concave supplementary sector developed between RP 1 and RP 2); basal hind margin not well preserved so that the presence of an anal angle or an anal triangle cannot be ascertained; no membranule visible. Discussion. The presence of a post-oblique-vein gap is a specialized character present in the Macromiidae and most taxa of Haplohamulida (sensu Bechly 2005). The presence of a well-defined Mspl, a well-developed, elongate, boot-shaped anal loop with a distinct toe and a distinct Cuspl) is a synapomorphy of the clade Italoansida Bechly, 1996 (Corduliidae Selys in Selys & Hagen, 1850, + Anauriculida Bechly, 1996). The following putative synapomorphies support a position in the Anauriculida (= Hemicorduliidae Bechly, 1996, + Libellulida Bechly, 1996) for Urolibellula gen. n.: two primary antenodal brackets Ax 1 and Ax 2 less distinct; besides anal crossing (= CuP) no cubito-anal crossveins present in hindwing (convergent to Pseudocordulia, Austrocordulia, Neophya, Hesperocordulia; maybe synapomorphic with some Corduliidae; reversed in some derived libellulids) because of reduction of the basal side of the hindwing subdiscoidal triangle; characteristic curved shape of Rspl that is distally ���rejoining��� IR 2; Mspl very distinct and curved, distally ���rejoining��� the MA. Urolibellula differs from the Hemicorduliidae (Hemicordulia Selys, 1870) in the most distal antenodal crossvein incomplete and oblique in forewing, presence of a one-celled subdiscoidal triangle in hindwing and discoidal triangle of forewing with three cells not separated by parallel crossveins. Urolibellula shares with the Libellulida (= Libellulidae Leach, 1815, + Urothemistidae Lieftinck, 1954) the following synapomorphies (note that in the phylogeny of the higher Libelluloidea proposed by Ware et al. (2007), the Urothemistidae are considered as a subfamily Urothemistinae of the Libellulidae but still falls as sister group of the remaining Libellulidae, in the same position as in Bechly (1996)): the two primary antenodal brackets Ax 1 and Ax 2 are very indistinct in forewings and indistinguishable from secondaries in hindwings (even if some secondary antenodals of forewing are not well aligned); anal loop reaching hind margin of hindwing or nearly so (well-developed ���toe���) (Figs. 4���5). Bechly (1996) proposed a further character for this clade, i.e. the second crossvein between RP 1 and RP 2 is developed as a ���libellulid oblique vein.��� This character is not very pronounced in Urolibellula, as this vein is only weakly oblique, but some modern Libellulidae (e.g. Libellula foliata Kirby, 1890) also have this vein not very oblique. Urolibellula differs from the Urothemistidae in the libellulid oblique vein not very oblique, most distal antenodal incomplete in forewing, presence of a subdiscoidal triangle in hindwing, and forewing discoidal triangle much broader (see Ris 1909 ��� 1913). Urolibellula shares with the Libellulidae the following apomorphies: most distal antenodal between costal margin and ScP developed as antenodal oblique vein, with suppression of corresponding antenodal between ScP and RA in forewing; all antenodal crossveins developed as antenodal brackets, therefore the two primaries Ax 1 and Ax 2 completely indistinguishable from the secondaries in hindwings. This character is also present in the forewings of many Libellulidae but not in all (e.g. Hydrobasileus Kirby, 1889, or the oldest libellulid Palaeolibellula). Nevertheless, Urolibellula lacks two other putative apomorphies of the Libellulidae, i.e. its sectors of arculus are not stalked and it has no apical supplement (strong concave supplementary sector developed between RP 1 and RP 2). Its forewing discoidal triangle is clearly broader than in any modern Libellulidae and divided into three smaller cells by crossveins not longitudinally disposed. Its subdiscoidal triangle and triangle are both very large with approximatively the same sizes. The modern Libellulidae with triangle and subdiscoidal triangle of similar sizes have a narrow triangle and a subdiscoidal triangle quite smaller than in Urolibellula. The forewing discoidal triangle and subdiscoidal triangle of Urolibellula reminds one of some ���corduliids��� (e.g. Epitheca Charpentier, 1825) (see Martin 1914), even of some Petalurida (Protolindenia Deichm��ller, 1886) or Aeshnoptera (Paramesuropetala Bechly et al., 2001) (Nel et al. 1998; Bechly et al. 2001). Thus this character is clearly subject to convergencies. In conclusion, the Urolibellulidae would fall in a ���basal��� position as probable sister group of the Libellulidae (sensu Bechly 1996, 2014), the Urothemistidae being the sister group of the (Urolibellulidae + Libellulidae). The presence of a representative of the crown group of the Libellulidae in the Late Cretaceous shows that the Urolibellulidae were certainly also present in the Late Mesozoic. The stem lineage of the Libellulidae was probably diversifying during the Late Cretaceous-Eocene even if the fossil record of this clade is very scarce for this period. Remarks. Cockerell (1921) described a fossil abdomen from the Eocene Green River Formation under the name Stenogomphus scudderi. The ���libelluloid��� genus Stenogomphus Scudder, 1892 is based on a forewing from the Earliest Oligocene Florissant Formation (Scudder 1892; Ris 1910; Nel & Paicheler 1994). There is no argument supporting the inclusion of S. scudderi in the genus Stenogomphus. S. carletoni Scudder, 1892 differs from our fossil in the complete last antenodal crossvein, and the second crossvein between RP 1 and RP 2 not oblique., Published as part of Zeiri, Asma, Nel, Andre & Garrouste, Romain, 2015, A new libelluloid family from the Eocene Green River Formation (Colorado, USA) (Odonata, Anisoptera), pp. 290-296 in Zootaxa 4032 (3) on pages 291-295, DOI: 10.11646/zootaxa.4032.3.4, http://zenodo.org/record/238343, {"references":["Bechly, G. (2005) A new fossil dragonfly (Anisoptera: Corduliidae) from the Paleocene Fur Formation (Mo clay) of Denmark. Stuttgarter Beitrage zur Naturkunde, (B), 358, 1 - 7.","Bechly, G. (1996) Morphologische Untersuchungen am Flugelgeader der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Berucksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Special Volume 2,1 - 402.","Ware, J. L., May, M. & Kjer, K. (2007) Phylogeny of the higher Libelluloidea (Anisoptera: Odonata): An exploration of the most speciose superfamily of dragonflies. Molecular Phylogenetic & Evolution, 45, 289 - 310. http: // dx. doi. org / 10.1016 / j. ympev. 2007.05.027","Ris, F. (1909 - 1913) Libellulinen monographisch bearbeitet. In: Collection Zoologique du Baron Edm. de Selys Longchamps. Catalogue systematique et descriptif, 9 - 16 bis, 1 - 1278.","Martin, R. (1914) Odonata. Fam. Libellulidae. Subfam. Cordulinae. Genera Insectorum, 155, 1 - 32.","Bechly, G., Nel, A., Martinez-Delclos, X., Jarzembowski, E. A., Coram, R., Martill, D., Fleck, G., Escuillie, F., Wisshak, M. M. & Maisch, M. (2001) A revision and phylogenetic study of Mesozoic Aeshnoptera, with description of several new families, genera and species (Insecta: Odonata: Anisoptera). Neue Palaontologische Abhandlungen, 4, 1 - 219.","Nel, A., Bechly, G., Jarzembowski, E. A. & Martinez-Delclos, X. (1998) A revision of the fossil petalurid dragonflies (Insecta: Odonata: Anisoptera: Petalurida). Paleontologia Lombarda, Milano, (N. S.), 10, 1 - 68.","Bechly, G. (2014) Phylogenetic Systematics of Odonata. Available from: http: // www. bernstein. naturkundemuseum-bw. de / odonata / intro. htm (accessed 14 October 2014)","Cockerell, T. D. A. (1921) Eocene insects from the Rocky Mountains. Proceedings of the United States National Museum, 57, 233 - 260. http: // dx. doi. org / 10.5479 / si. 00963801.57 - 2313.233","Scudder, S. H. (1892) Some insects of special interest from Florissant, Colorado and other points of the Tertiaries of Colorado and Utah. Bulletin of the United States Geological Survey, 93, 25 pp.","Ris, F. (1910) The identity of two Odonata fossils. Bulletin of the Wisconsin Natural History Society, Milwaukee, 8, 102 - 105.","Nel, A. & Paicheler, J. - C. (1994) Les Libelluloidea fossiles autres que Libellulidae. Un inventaire critique (Odonata, Corduliidae, Macromiidae, Synthemistidae, Chlorogomphidae et Mesophlebiidae). Nouvelle Revue d'Entomologie, (N. S.), 11, 321 - 334."]}

Araripescolia magnifica Nel, Escuillie & Garrouste, sp. nov. (Figs 1���3) Material. Holotype specimen WDC-CCFB- 7 (a complete body with four wings and two legs), Wyoming Dinosaur Center-Crato, Wyoming, USA. Etymology. Named after the wonderful state of preservation of the wings of this fossil. Age and outcrop. Chapada do Araripe, northeastern Brazil; Upper Aptian, Nova Olinda member of the Crato formation. Diagnosis. As for the genus. Description. Head length 4.2 mm, 4.1 mm wide, mesosoma 7.4 mm long, 7.1 mm wide, metasoma 12.9 mm long, 5.8 mm wide; fore wing 16.1 mm long, 5.8 mm wide; hind wing 10.9 mm long, 4.5 mm wide; head mediumsized and suboval; eyes large but poorly preserved; details of head structures not clear; antennae long, curved, with separation between distal segments indistinct; scape enlarged; pedicel small; mesosoma large; pronotum short medially; scutellum inversely trapezoidal; metanotum short; propodeum broad; wings hyaline, presence of longitudinally wrinkled structures of apical parts of fore and hind wings; fore wing with pterostigma narrow, widened apicad, slightly sclerotised; costal vein confluent before pterostigma; radial vein uninterrupted; basal section of Rs as long as that of M; no vestigial 1 r-rs; distal section of Rs sigmoidal, 2.7 mm long; crossvein r-rs short, straight; veins 3 r-m and 2 m-cu present; veins 1 m-cu and 2 m-cu ending in cell 1 + 2 Rs; anterior side of cell 1 + 2 Rs 0.5 mm long; cell 2 R 1 1.5 times as long as pterostigma, three times as long as wide, with apex not contiguous with wing margin; cell 3 Rs closed, nearly as long as wide; cu-a aligned with basal section of M, slightly shorter than basal section of M; cell 2 Cu nearly parallel-sided, with sides arcuate; hind wing with Rs short beyond 1 r-m; 1 r-m long and curved; cell 1 Rs closed, longer than wide; cu-a postfurcal, long sigmoidal and oblique to A; metasoma long (longer than head and thorax combined), somewhat constricted between 1 st and 2 nd segments; sting not clearly visible. Discussion. Araripescolia gen. nov. has a fore wing venation very similar to that of an Archaeoscoliinae (very similar to that of the Mesozoic genera Archaeoscolia Rasnitsyn, 1993 or Cretoscolia Rasnitsyn, 1993) (Rasnitsyn 1993), in the following points: fore wing cell 2 R 1 long; cell 3 Rs closed; crossvein r-rs short, straight; crossveins 3 rm and 2 m-cu present; hind wing Rs and M very long before r-m. Cretoscolia brasiliensis is the unique described Archaeoscoliinae from Araripe, it is much smaller than Araripescolia, and it has not the longitudinally wrinkled structure of the apical parts of the fore wings, clearly visible in Araripescolia (Osten 2007). Rasnitsyn (1993) considered the Archaeoscoliinae as a paraphyletic group in respect to the other Scoliidae. Argaman (1996) considered that the Archaeoscoliinae are not Scoliidae but ���Anthoboscidae��� (= Tiphiidae: Anthoboscinae) for the presence of notauli. This structure is not preserved in Araripescolia. Later authors did not follow Argaman���s opinion (Rasnitsyn & Mart��nez-Delcl��s 1999, Zhang et al. 2002). After Pilgrim et al. (2008), the Tiphiidae is a polyphyletic group with the Anthoboscinae clearly not closely related to the Scoliidae, while the Scoliidae would belong to a clade that comprises the Formicidae and Apoidea. Notauli are present in these two last groups, suggesting that their absence in modern Scoliidae is an apomorphy. The plesiomorphic presence of notauli in the Archaeoscoliinae is therefore not a sufficient argument to exclude them from the scoliid lineage. It remains that a phylogenetic analysis of the Scoliidae that integrates these fossils is necessary to clarify the situation. Some Anthoboscinae (e.g. Tiphiodes Br��thes, 1913) have a fore wing venation very similar to those of Araripescolia and Cretoscolia (Genise 1984). Araripescolia can be attributed to the Scoliidae because of the presence of the typical longitudinally wrinkled structures of the apical parts of the fore and hind wings, absent in the Tiphiidae (Day et al. 1981, Argaman 1996). These structures are also absent in the archaeoscoliine genera Cretoscolia and Archaeoscolia (Rasnitsyn 1993, Rasnitsyn & Mart��nez-Delcl��s 1999, Zhang 2004, Osten 2007), and badly known in the two other archaeoscoliine genera, viz. the Mesozoic Protoscolia Zhang et al., 2002 and the Cenozoic Floriscolia Rasnitsyn, 1993 (Rasnitsyn 1993, Zhang et al. 2002). Araripescolia strongly differs from Cenozoic Floriscolia Rasnitsyn, 1993 in the fore wing with apex of cell 2 R 1 not contiguous with wing margin, cell 3 Rs nearly as long as high, and hind wing with cu-a well distal of M-Cu fork (Rasnitsyn 1993). Araripescolia differs from Protoscolia Zhang et al., 2002 in the absence of a vestige of 1 r-rs, and the apex of radial cell nearly aligned to 3 r-m (Zhang et al. 2002). Thus Araripescolia can be accurately attributed to the Scoliidae, but, if it strongly differs from the Archaeoscoliinae, its exact relationships with this group remain uncertain. Araripescolia shares with the Scoliinae (sensu Rasnitsyn 1993, including the Campsomerinae and Colpinae sensu Argaman 1996) the following characters: the costal vein is confluent before the pterostigma, while it is interrupted in the Proscoliinae and in Archaeoscoliinae (Rasnitsyn, pers. comm.); the radial vein is uninterrupted in the Scoliinae, while it is very narrow at the pterostigmal base in Proscoliinae and almost interrupted in some Archaeoscoliinae (Day et al. 1981, Rasnitsyn & Mart��nez-Delcl��s 1999). The character ���interruptions of costal and radial veins��� is the general ���rule��� in the clade that comprises the Scoliidae, Formicidae and Apoidea. Thus the absence of interruption would rather be an apomorphic character, supporting a position of Araripescolia in the Scoliinae sensu Rasnitsyn (1993) and a possible basal position for the Proscoliinae. Araripescolia differs from the Campsomerinae Betrem, 1972 (sensu Argaman 1996) (a group reduced to the two genera Carbonelis Betrem, 1972 and Dasyscolia Bradley, 1951), in the presence of a fore wing cell 1 + 2 Rs with a anterior side long and the presence of a closed cell 3 Rs. Araripescolia differs from the modern Scoliinae (sensu Argaman, 1996) in the presence of two veins ending in cell 1 + 2 Rs and therefore it would rather fall near the Colpinae Argaman, 1996. Some Colpinae have long closed cells 2 R 1 and 3 Rs as in Araripescolia but they generally differ from Araripescolia in their cell 1 + 2 Rs with anterior side much shorter (or reduced to a point) (Betrem 1972, Argaman 1996). Unfortunately the body structures of Araripescolia are not enough well preserved to be compared to those of the modern Scoliidae. The hind wing cell 1 Rs closed of Araripescolia is quite a rare character in Apocrita and can be considered as an autapomorphy of this taxon (Rasnitsyn, pers. comm.) Most modern scoliid species are external parasitoids of soil-inhabiting scarab beetle larvae. The Scarabaeidae are well known in the Early Cretaceous, and dung beetles are recorded from the Crato formation (Wolf- Schwenninger & Schawaller 2007) suggesting a similar biology for these Cretaceous Scoliidae. Modern scoliid adults are frequently floricolous and could participate to pollination, but it is delicate to infer such behaviours for the Early Cretaceous., Published as part of Nel, Andr��, Escuillie, Fran��ois & Garrouste, Romain, 2013, A new scoliid wasp in the Early Cretaceous Crato Formation in Brazil (Hymenoptera: Scoliidae), pp. 395-400 in Zootaxa 3717 (3) on pages 396-398, DOI: 10.11646/zootaxa.3717.3.10, http://zenodo.org/record/221964

Nel, André, Escuillie, François, Garrouste, Romain (2013): A new scoliid wasp in the Early Cretaceous Crato Formation in Brazil (Hymenoptera: Scoliidae). Zootaxa 3717 (3): 395-400, DOI: http://dx.doi.org/10.11646/zootaxa.3717.3.10

Genus Araripescolia Nel, Escuillie & Garrouste, gen. nov. Type species. Araripescolia magnifica Nel, Escuillie & Garrouste, sp. nov. Etymology. Named after the type locality Sierra of Araripe and the genus name Scolia, gender feminine. Diagnosis. Presence of longitudinally wrinkled structures of apical parts of fore and hind wings; fore wing cell 2 R 1 long, with apex not contiguous with wing margin; cell 3 Rs closed, nearly as long as high; cell 1 + 2 Rs with a long anterior side; crossvein r-rs short, straight; crossvein 2 m-cu sigmoidal; costal vein confluent before pterostigma; radial vein uninterrupted; hind wing Rs and M very long before r-m; cell 1 Rs closed; cu-a well distal of M-Cu fork., Published as part of Nel, Andr��, Escuillie, Fran��ois & Garrouste, Romain, 2013, A new scoliid wasp in the Early Cretaceous Crato Formation in Brazil (Hymenoptera: Scoliidae), pp. 395-400 in Zootaxa 3717 (3) on page 396, DOI: 10.11646/zootaxa.3717.3.10, http://zenodo.org/record/221964

Family Scoliidae Latreille, 1802, Published as part of Nel, Andr��, Escuillie, Fran��ois & Garrouste, Romain, 2013, A new scoliid wasp in the Early Cretaceous Crato Formation in Brazil (Hymenoptera: Scoliidae), pp. 395-400 in Zootaxa 3717 (3) on page 396, DOI: 10.11646/zootaxa.3717.3.10, http://zenodo.org/record/221964

Genus Isometopsallops Herczek & Popov, 1992 Type species. Isometopsallops schuhi Herczek & Popov, 1992., Published as part of Vernoux, Julien, Garrouste, Romain & Nel, Andre, 2010, The first psallopinous bug from Lowermost Eocene French amber (Hemiptera: Heteroptera: Miridae), pp. 63-68 in Zootaxa 2499 on page 63, DOI: 10.5281/zenodo.195786, {"references":["Herczek, A. & Popov, Yu. A. (1992) A remarkable psallopinous bug from Baltic amber (Miridae). Mitteilungen der Geologische-Palaontologische Institute der Universitat Hamburg, 73, 235 - 239."]}

Isometopsallops prokopi sp. nov. Figs 1���4 Material. Holotype MNHN A 30851 (PA 8543, male), allotype MNHN A 30852 (PA 6524, female), paratypes MNHN A 30853 (PA 8584, male), and MNHN A 30854 (PA 9493, male), all stored at the Laboratoire de Pal��ontologie, Mus��um national d���Histoire naturelle, Paris, France. Etymology. Named after our friend and colleague Dr Jakub Prokop (Charles University, Praha). Diagnosis. tarsi three-segmented; a subapical tooth on each claw; parempodia setiform; no pulvilli; broad femora; seven metafemoral trichobothria and six mesofemoral trichobothria; large eyes semi-globular; antennae placed basally on inner side of eyes; head nearly vertical; no distinct collar; pronotum trapezoidal, ratio length/width pronotum 0.4; membrane of hemelytra crumpled and with two cells, one small and one large; body two times as long as wide, with dorsal surface smooth, with dense hairs; male right genital style (or paramere) slightly longer than left one. Description. General coloration light brown or brownish; macropterous; dorsal surface smooth, impunctate, with moderately short, erect and sub-erect hairs, densely covering dorsal surface of body and posteriorly directed. Head large, nearly spherical in frontal view, nearly vertical, and nearly twice as wide as long; antenna cylindrical, inserted at inner side of eyes about 1 / 3 of their lower level; eyes very large, almost semi-globular, covering much of dorsal surface of head, bare, emarginated along inner side of genae; interocular space much narrower than width of eye, hind margins of eyes touching anterior margin of pronotum; mandibular plates not enlarged; labium four-segmented, inserted ventrally on head, and reaching at least a third of abdomen, with segment 1 reaching posterior margin of head; trapezoidal pronotum a little longer than head, with ratio length/width = 0.4, collar absent; no calli; lateral margins and posterior margin not carinate nor emarginated; mesoscutum two times shorter than scutellum; claval commissure present; cuneus and costal fracture clearly distinct; a white spot or light-coloured spot around the costal fracture; hemelytral membrane with a large cell and a small cell which branches to a supplementary vein, membrane distinctly crumpled, hairs absent on membrane; femora of hind legs broad and long, tibia nearly as long as femora, seven metafemoral trichobothria and six mesofemoral trichobothria; tibia with delicate spines distally, three-segmented tarsi more or less of equal size, first tarsal segment well separated from second but suture between third and second segments is weaker than that between first and second (probably presence of a pseudo-trisegmentation sensu Gorczyca [2000]); claws of equal size and almost straight; a subapical tooth on each claw; parempodia setiform; no pulvilli. Dimension. (taken on PA 8543 and PA 9493 for accessibility): Length of body from apex of hemelytra 2.25 mm (around 2.5 mm for the PA 8584 and PA 6524); width 1.025 mm; length of hemelytra 1.75 mm; length of head between 0.25 and 0.375 mm (for different bending of head); width of head 0.625 mm; height of head 0.375 mm; width of eye 0.3 mm; distance between eyes 0.075 mm; antennal segments IV 0.225 mm long, III 0.275 mm long, II 0.4 mm, I 0.115 mm, antennal segment II the longest; length of pronotum 0.35 mm, width 0.875 mm (max.), 0.375 mm (min.); length of mesoscutum 0.15 mm; length of scutellum 0.27 mm; claval commissure 0.525 mm long; costal fracture 0.25 mm long; hind femora 0.87 mm long, (1.05 mm for PA 6524), 0.25 mm wide; hind tibia 0.85 mm long, width mm, tarsal joints I 0.083 mm, II+III 0.180 mm long. Discussion. After Herczek & Popov (1992) and Popov & Herczek (2006), the mirid subfamily Psallopinae is characterized by the following features: large eyes covering greater part of head, very deeply inserted antennae, absence of ocelli, and presence of subapical tooth on claws. These authors placed the two genera Isometopsallops and Cylapopsallops in this subfamily on the basis of these characters. As they are also shared by Isometopsallops prokopi sp. nov., we consider that it also belongs to this subfamily. One further character that potentially supports an attribution of I. prokopi to the Psallopinae is the presence of seven metafemoral trichobothria, a putative apomorphy also present in the Phylinae, after Schuh (1976: 13). The Isometopinae, Cylapinae, and Psallopinae share also plesiomorphic characters, i.e., parempodia setiform (Schuh & Schwartz, 1984; Schuh & ��tys, 1991); and absence of pulvilli (Schuh [1976] observed no pulvilli in Isometopinae, Psallopinae, and Phyllinae; for Cylapinae, pulvilli are usually present and attached to ventral surface of claw which is an apomorphic character); antennae in inner side of eyes (Schuh and Schwartz, 1984; Schuh and ��tys, 1991). The Psallopinae is currently considered to be related to the two subfamilies, Cylapinae and Isometopinae (Gorczyca, 2000; Lin & Yang, 2005). Gorczyca (2000) supported the clade (Cylapinae + (Psallopinae + Isometopinae)) on the basis of two putative synapomorphies: presence of claws with a subapical tooth, and tarsi two-segmented. The presence of three-segmented tarsi in I. prokopi, I. schuhi, and Cylapopsallops would then be problematic for their attribution to the Psallopinae. Schuh & Schwartz (1984) and Schuh & ��tys (1991) considered three-segmented tarsi as plesiomorphic. This character state is present in other mirid subfamilies. However, there are some cylapine genera as Yamatofulvius where all the species have apparently functional three-segmented tarsi (Yasunaga, 2000). Schuh (1976) described also Cylapinae and Isometopinae with tarsi two- or three-segmented (for Isometopinae, see also Herczek [1993]). Thus this character is at least homoplasic. Lastly, Lin & Yang (2005) indicated: "according to Schuh & Slater (1996: 172) tarsi are twosegmented in the Isometopinae and Psallopinae; two- or three-segmented in the Cylapinae; and threesegmented in the Orthotylinae, Phylinae, Bryocorinae, Deraeocorinae, and Mirinae." The two-segmented tarsus is judged to be plesiomorphic by these last authors. The presence of large eyes (also present in the three fossils I. prokopi, I. schuhi, and Cylapopsallops) would be the main apomorphic characters supporting the clade (Isometopinae + Psallopinae) (Schuh & Schwartz, 1984; Schuh & ��tys, 1991; Gorczyca, 2000). Within the clade (Isometopinae + Psallopinae), the absence of ocelli characterises the Psallopinae and is the unique apomorphy visible on the three fossils I. prokopi, I. schuhi, and Cylapopsallops. For Herczek (1993), it seems obvious that the occurrence of ocelli is plesiomorphic. Schuh & Schwartz (1984) preferred the interpretation of character distributions as a convergent lose of the ocelli in Psallopinae and Cylapinae. I. prokopi differs from the recent Psallops in that the right genital style is larger than the left one (Yasunaga, 1999; Lin, 2004; Lin & Yang, 2005). In recent Cylapinae, the genital styles are asymmetrical in shape and size, one either left or right larger, depending on the species or genus. Thus nearly all the characters currently used to support the Psallopinae, Cylapinae, and Isometopinae relationships are homoplastic. And so the positions of the three fossils (I. prokopi, I. schuhi, and Cylapopsallops) may perhaps be subject to changes in the future. I. prokopi differs from Cylapopsallops because its body is less elongate (only two times as long as wide, not three times as long as wide); broad femora, not flattened in Cylapopsallops; collar absent, instead of being distinct in Cylapopsallops. The habitus of I. prokopi is very similar to that of I. schuhi. They share the same characters mentioned above: tarsi three-segmented (apparently pseudo-trisegmentation); subapical tooth on each claw; trapezoidal pronotum; and two-celled membrane of hemelytra. The ratio length/width of the pronotum = 0.4 in I. prokopi but 0.66 in Isometopsallops schuhi. We lack information on the number of femoral trichobothria in the latter, which renders their comparison for now a little less positive. With the present state of knowledge we prefer to include our fossil in a new species of the same genus Isometopsallops. Unfortunately the male genitalia of I. schuhi are not described. The presence of a new species belonging to the Baltic amber genus Isometopsallops in the Lowermost Eocene French amber is not surprising because similar fossils occur in other orders (Psocoptera, etc.) (Nel et al., 2005). The genus Psallops comprises an undescribed Eocene species from Baltic amber, plus 10 known recent species in the world (Schuh, 1995; Yasunaga, 1999; Lin, 2004; Popov & Herczek, 2006): P. grandoculus Linnavuori & Alamy, 1982 from Saudi Arabia; P. oculatus Usinger, 1946 from Guam and the Mariana Islands, Saipan; P. ponapensis Carvalho, 1956 and P. yapensis Carvalho, 1956 from the Caroline Islands; P. myiocephalus Yasunaga, P. nakatanii Yasunaga, 1999, and P. yaeyamanus Yasunaga, 1999 from Japan; P. formosanus Lin, 2004 and P. l e e a e Lin, 2004 from Taiwan; and P. c h i n e n s i s Lin, 2004 from China. The second recent psallopsine genus Isometocoris is from Panama. Thus the recent Psallopsinae live under very different climates even if all are warm. One would infer a warm palaeoclimate for the three fossils I. prokopi, I. schuhi, and Cylapopsallops. But as we know nothing about the exact phylogenetic relationships between them and the recent Psallopsinae, and also because the recent representatives of the Isometopinae (putative psallopsine sister group) can be found in very diverse environments and climate, in Europe, North America, Africa, etc., it is in fact not possible to infer anything on the biology of these fossil taxa after their taxonomic position (Nel, 1997). Nevertheless, recent Psallopinae seem to live on tree trunks and bark (Yasunaga, 1999). If the corticolous habits may also be inferred for the Eocene Psallopinae, it would not be a surprise to have an overrepresentation of this subfamily in amber inclusions., Published as part of Vernoux, Julien, Garrouste, Romain & Nel, Andre, 2010, The first psallopinous bug from Lowermost Eocene French amber (Hemiptera: Heteroptera: Miridae), pp. 63-68 in Zootaxa 2499 on pages 63-68, DOI: 10.5281/zenodo.195786, {"references":["Gorczyca, J. (2000) A systematic study on Cylapinae with a revision of the Afrotropical region (Heteroptera, Miridae). Wydawnictwo Uniwersystetu Slaskiego, Katowice, 176 pp.","Herczek, A. & Popov, Yu. A. (1992) A remarkable psallopinous bug from Baltic amber (Miridae). Mitteilungen der Geologische-Palaontologische Institute der Universitat Hamburg, 73, 235 - 239.","Popov, Yu. A. & Herczek, A. (2006) Cylapopsallops kerhneri gen. et sp. n. - a new peculiar mirid from Baltic amber (Heteroptera: Miridae: Psallopinae). Russian Entomological Journal, 15 (2), 187 - 188.","Schuh, R. T. (1976) Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of relationships within the family. American Museum Novitates, 2601, 1 - 26.","Schuh, R. T. & Schwartz, M. D. (1984) Carvalhoma (Hemiptera: Miridae): revised subfamily placement. Journal of the New York Entomological Society, 92, 48 - 52.","Schuh, R. T. & Stys, P. 1991. Phylogenetic analysis of cimicomorphan family relationships (Heteroptera). Journal of the New York Entomological Society, 99, 298 - 350.","Lin, C. - S. & Yang, C. - T. (2005) External male genitalia of the Miridae (Hemiptera: Heteroptera). National Museum of Natural Science Special Publication, 9, 1 - 174.","Yasunaga, T. (2000) The mirid subfamily Cylapinae (Heteroptera: Miridae) or fungal inhabiting plant bugs in Japan. Tijdschrift voor Entomologie, 143, 183 - 209.","Herczek, A. (1993) Systematic position of the Isometopinae Fieb. (Miridae, Heteroptera) and their interrelationships. Prace Naukowe Uniwersytetu S l q skiego, 1357, 1 - 86.","Yasunaga, T. (1999) First record of the plant bug subfamily Psallopinae (Heteroptera: Miridae) from Japan, with descriptions of three new species of the genus Psallops Usinger. Proceedings of the Entomological Society of Washington, 101, 737 - 741.","Lin, C. - S. (2004) First record of the plant bug subfamily Psallopinae (Heteroptera: Miridae) from Taiwan and China, with descriptions of three new species of the genus Psallops Usinger. Formosan Entomology, 24, 273 - 279.","Nel, A., Prokop, J., De Ploeg, G. & Millet, J. (2005) New Psocoptera (Insecta) from the Lowermost Eocene amber of Oise, France. Journal of Systematic Palaeontology, 3, 371 - 391.","Schuh, R. T. (1995) Plant bugs of the World (Insecta: Heteroptera: Miridae): systematic catalog, distributions, host list, and bibliography. The New York Entomological Society, New York, 1329 pp.","Nel, A. (1997) The probabilistic inference of unknown data in phylogenetic analysis. In: Grancolas, P. (ed.), The origin of biodiversity in insects: phylogenetic tests of evolutionary scenarios. Memoires du Museum National d'Histoire Naturelle, Paris, 173, 305 - 327."]}

Zeiri, Asma, Nel, Andre, Garrouste, Romain (2015): A new libelluloid family from the Eocene Green River Formation (Colorado, USA) (Odonata, Anisoptera). Zootaxa 4032 (3): 290-296, DOI: http://dx.doi.org/10.11646/zootaxa.4032.3.4