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Possible Furagrion FUM-N 17242 (Fig. 24): poorly preserved partial head, thorax, part of the abdomen and four legs, indistinct basal parts of all wings, coll. Jan and Elly Verkleij, 2010, Skarrehage, Mors, diatomite, near ash layer -13, in the Fur Museum. Resembles, but might not be Furagrion., Published as part of Archibald, S. Bruce, Ware, Jessica L., Rasmussen, Jan A., Sylvestersen, René L., Olsen, Kent & Simonsen, Thomas J., 2023, The damselfly genus Furagrion Petrulevičius et al. (Odonata, Zygoptera) from the early Eocene Fur Formation of Denmark and the dysagrionoid grade, pp. 289-317 in Zootaxa 5278 (2) on page 314, DOI: 10.11646/zootaxa.5278.2.4, http://zenodo.org/record/7906172

Genus Furagrion Petrulevičius, Wappler, Wedmann, Rust,and Nel,2008: 176. Type species: Phenacolestes jutlandica Henriksen, 1922, by original designation, Published as part of Archibald, S. Bruce, Ware, Jessica L., Rasmussen, Jan A., Sylvestersen, René L., Olsen, Kent & Simonsen, Thomas J., 2023, The damselfly genus Furagrion Petrulevičius et al. (Odonata, Zygoptera) from the early Eocene Fur Formation of Denmark and the dysagrionoid grade, pp. 289-317 in Zootaxa 5278 (2) on page 292, DOI: 10.11646/zootaxa.5278.2.4, http://zenodo.org/record/7906172, {"references":["Petrulevicius, J. F., Wappler, T., Wedmann, S., Rust, J. & Nel, A. (2008) New Megapodagrionid Damselflies (Odonata: Zygoptera) from the Paleogene of Europe. Journal of Paleontology, 82, 1173 - 1181.","Henriksen, K. (1922) Eocene insects from Denmark. Danmarks Grologiske UndersOgelser, II Raekke, 37, 1 - 36."]}

Furagrion ansorgei (Zessin, 2011) comb. nov. Figs. 22–23 Emended diagnosis. The wing of Furagrion ansorgei may be distinguished from those of F. jutlandicus as in its diagnosis, above. Emended description. Holotype: see genus description and its distinctive character states in the diagnosis of F. jutlandicus. Type material. Holotype (Fig. 23) MOA 770/1, 2: likely a forewing by its slender shape (Fig. 23B), coll. Jörg Ansorge, northern coast of Mors, Skaerbaek, Fur Formation, in a calcareous concretion, in the Natural History Museum, Natural Research Society Mecklenburg, Natureum at Ludwigslust Castle (part) and Natural History Museum of Denmark, Copenhagen, Denmark (counterpart). We were not able to examine the fossil as the part and counterpart could not be located at the institutions listed. Our discussion is based on Zessin’s (2011) description and illustrations, and a higher resolution copy of his figure 3 provided by Jörg Ansorge. Other material. MGUH 34113 (Fig. 22) tentatively belongs to the species: a complete forewing in a concretion, coll. Thomas Klode on Fur Island, Knudeklint Member, possibly between ash layers -11 and -13, Fur Formation, 1968, in the Natural History Museum of Denmark, University of Copenhagen. Range and age. Earliest Ypresian Fur Formation, Jutland, Denmark. Range: see type and other material, above. Remarks. With the ability to examine a larger sample of Furagrion specimens than was available to Zessin (2011), we find that the character states of Morsagrion that he proposed would separate it from Furagrion do not do so. These are: 1- The RA-RP1 space is two cells wide from about the middle of the level of the pterostigma. - We find this difficult to discern from his fig. 3 photograph and we are not confident that this is established. 2- The form of the pterostigma. - We find that it does not significantly differ. 3- The relationship between Zessin’s “premedial cell” (“prmc”) and the cells below it in the MA-MP space. The premedial cell is the space between RP to slightly past the base of RP3-4 and MA without crossveins from the arculus to the first crossvein between RP-3-4 and MA. He excludes the quadrangle and counts four cells in the MA-MP space subtending this in Morsagrion and two cells in this space in Furagrion. - By our count, this is about three and a half cells long in M. ansorgei, and in the larger sample of Furagrion available to us, two (e.g., FUM-N 13856, Fig. 2) to four (FUM-N 14704, Fig. 17) cells long and, therefore, consider this to be within intraspecific variation. 4- The number of cells in the precubital field (MP-CuA space) to the level of the origin of RP3-4 (three in Morsagrion n. gen., four at Furagrion). - Like Zessin, we count three in M. ansorgei, but with our larger sample we find between two (hind wings of MM-10752) to five (FUM-N 11146) in Furagrion. We, therefore, also consider this to be intraspecific variation. 5- Eight cells between the beginning of IR2 and RP 2 in Morsagrion and five in Furagrion. This varies from eight cells in Henriksen’s type MGUH 1819 to five in FUM-N 13856, with fewer in hind wings, e.g., MM-10750 with 7.5 and 6.5 cells in the forewings and 4.5 in the hind wing. - In all of these except MGUH 34113, which we tentatively associate with MOA 770, the IR1-RP2 space is more than one cells wide distad about the level of the pterostigma base (see below). 6- The shape of the discoidal and subdiscoidal cell. - We find that this does not greatly differ. 7- The length of the double rows of cells in the IR1-RP2 space. - This space is more than one cell wide for only about three cells from ending on the wing margin, whereas in F. jutlandicus, this space is more than one cell wide to about the level of or slightly more basal of the basal end of the pterostigma. This occurs in all Furagrion specimens that we examined except possibly MGUH 34113 (below) and does appear distinctive. We, therefore, find that MOA 770/1and MGUH 34113 do not differ enough from the fossils assigned here to F. jutlandicus to justify placing them in a separate genus, and so treat Morsagrion as a junior synonym of Furagrion. Character state 7 of MOA 770 does appear distinctive among the Furagrion specimens examined (but see MGUH 34113, below), supporting its status as a separate species. However, while it is shown in Zessin’s drawing (his figs. 1 and 4), this region appears unclear in his photograph (his fig. 3) and in a slightly higher resolution version of it provided by J. Ansorge (see our drawing made from this, Fig. 23). We consider this character state as tentative until the type specimen can be examined. The shape of wing MOA 770 provides stronger support for recognizing it as a separate species (Fig. 23B, 23C). It has a greater length / width ratio than all Furagrion wings, fore- or hind, by all length measurements except RP2 to apex / width (Tables 2 and 3 and see PCA, above). By these reasons, we treat MOA 770 as Furagrion ansorgei (Zessin) comb. nov. MGUH 34113 (Fig. 22) might be conspecific with MOA 770: Its IR1-RP2 space becomes more than one cell wide closer to the level of the distal end of pterostigma than basal, suggesting an association with F. ansorgei MOA 770. This is, however, not as distinct as in Zessin’s drawing of the MOA 770 wing. More importantly, this wing is relatively narrow like MOA 770. Furthermore, it is smaller (see PCA and Tables 3 and 4), although this might represent intraspecific variation or sexual dimorphism. Given this uncertainty, we tentatively treat this specimen as F. ansorgei., Published as part of Archibald, S. Bruce, Ware, Jessica L., Rasmussen, Jan A., Sylvestersen, René L., Olsen, Kent & Simonsen, Thomas J., 2023, The damselfly genus Furagrion Petrulevičius et al. (Odonata, Zygoptera) from the early Eocene Fur Formation of Denmark and the dysagrionoid grade, pp. 289-317 in Zootaxa 5278 (2) on pages 312-314, DOI: 10.11646/zootaxa.5278.2.4, http://zenodo.org/record/7906172, {"references":["Zessin, W. (2011) Neue Insekten aus dem Moler (Palaozan / Eozan) von Danemark Teil 1 (Odonata: Epallagidae, Megapodagrioniidae), Virgo, Mitteilungsblatt des Entomologischen Vereins Mecklenburg, 14, 64 - 73."]}

Morsagrion Zessin, 2011: 65. Type species: Morsagrion ansorgei Zessin, 2011, by original designation. Syn. nov. Figs. 1–4, 6–8, 1124. Included species. Furagrion jutlandica (Henriksen, 1922), Furagrion ansorgei (Zessin, 2011) comb. nov. Range and age. Earliest Ypresian Fur Formation, Jutland, Denmark. Specimens of this genus are known to range from just below ash layer -13 within the Knudeklint Member to the lower part of the Silstrup Member (the ash layer +25 – +30 interval) of the Fur Formation, which has an absolute age of approximately 55.5 Ma (Storey 2007, Stokke et al. 2020). Six of the sixteen examined fossils, however, are from unknown levels within the Fur Formation. Emended diagnosis. The wings of Furagrion differ from those of similar extant and extinct Zygoptera by possession of all nine wing character states of the Dysagrionidae Cockrerell (Cephalozygoptera) diagnosis listed below, but the genus is excluded from that family and suborder by their zygopteran bulging, hemispherical compound eyes set far apart on a short head (see Archibald et al. 2021). Wings most easily distinguished from those of other Zygoptera and from taxa that possess many of the nine wing character states listed below and might be Zygoptera or Cephalozygoptera as follows: from Viridiflumineagrion Nel by pterostigma ca. 2.5 times longer than wide [ca. 4.7 times], from Miopodagrion Kennedy by C-RA space distad pterostigma with one row of cells [apparently two full rows]. Wings distinguished from genera with similar wings without an associated head by one or more of the following: 10, C-RA space distad pterostigma almost always with a single row of cells, rarely a few rows two cells wide immediately distad pterostigma; 11, CuA-A space usually two, sometimes maximum three cells wide; 12, possession of a brace vein; 13, no accessory (“secondary”) antenodal crossveins: Allenbya Archibald and Cannings: [10: 2 and 3 cells wide throughout]; Valerea Garrouste et al.: [10: many, up to five cells wide]; Thanetophilosina Nel et al.: [12: no brace vein]; Electrophenacolestes Nel and Arillo: [11: up to five wide, 13]; Menatagrion Nel and Jouault, 2022 [12, 13]; Chickaloon specimen of Garrouste & Nel (2019): [11: up to four wide]; Specimen MeI6572 (Megapodagrionidae genus and species A of Petrulevičius et al. 2008, cf. Megapodagrionidae genus and species A of Archibald et al. 2021): [10: most of C-RA space two cells wide]. Emended description. Head short. Compound eyes bulging, hemispherical, set apart twice their width in dorsal aspect. Thorax, legs generalised as known by preservation. Forewing. Membrane darkly infuscate, but some might be hyaline as preserved (see below). Measurements, ratios of these see Tables 1 and 2. Antenodal crossvein Ax0 absent, Ax1 and Ax2 somewhat lengthened to moderately expand antenodal space. No accessory antenodal crossveins. Pterostigma ca. 2–3 times longer than wide; anterior, posterior margins oblique; subtends 2–3 cells; distinctly oblique brace vein at basal-posterior corner in RA-RP1 space. Crossveins in postnodal, postsubnodal spaces mostly aligned basally, usually not distally. C-RA space distad pterostigma almost always with a single row of cells, rarely briefly two cells wide (see F. jutlandicus proposed neotype, Figs. 3–4); RA meets margin at or very near apex; slightly upturned near margin. Wing dense with cells throughout. RP2: origin 6.5–9 cells distal to origin of IR2. IR2: origin at or very near, basal to subnodus. RP3-4: origin about 3/5 from arculus to subnodus. Arculus at or very close basad Ax2. All major veins linear except MA zigzagged distad ca. mid-way between arculus, termination; CuA zigzagged, slightly near quadrangle, increasingly toward terminus. No crossvein O. CuA terminates on posterior margin ca. mid-way between nodus, apex; CuA–A space 2–3 cells wide at widest. Hind wing. Like forewing except shorter relative to width; measurements, ratios of these see Tables 1 and 2. Abdomen generalised as known by preservation. Remarks. Proposed neotype designation. Henriksen’s (1922) holotype and only specimen then known (Fig. 1) is incomplete, consisting of two wings that are partially complete distal to the nodus, the mid-posterior fragment of a third, and an abdomen that is complete except for a portion of its base (MGUH 1819, Natural History Museum of Denmark, University of Copenhagen, Denmark). Lacking information from the diagnostically important antenodal region of the wings, he assigned the species to Phenacolestes, an extinct genus of the extinct suborder Cephalozygoptera known from the Eocene and possibly Miocene (see table 3 of Archibald et al. 2021) with similar venation in its preserved portions. Cockerell (1908) had assigned the genus to the Dysagrioninae, then a zygopteran subfamily of Agrionidae Leach (now Calopterygidae Sélys). Recognising the limitations of the incomplete type specimen, Nel & Paicheler (1994) considered the species ‘ Phenacolestes ’ jutlandica as family indet. Rust (1999) expressed even less confidence in the generic assignment, treating the species as ‘Dysagrioninae gen. indet. jutlandica ’, illustrating it (his fig. 4 and plate 1, fig. a) with the non-type specimen FUM-N 13856 (then ERK KL Tl), an almost complete isolated wing and 14M-A2163 (his plate 1, fig. b). Petrulevičius et al. (2008) assigned the species to their new, monotypic genus Furagrion based on ERK-KL-T1 (FUM-N 13856) (Fig. 2). Zessin (2011) subsequently described Furagrion morsi and Morsagrion ansorgei, erroneously citing FUM-N 13856 (as ERK-KL-T1) of Petrulevičius et al. as the holotype of Furagrion jutlandicus. Archibald et al. (2021) considered Furagrion a dysagrionid in the Cephalozygoptera. We agree that FUM-N 13856 is conspecific with Henriksen’s holotype by the extensive similarity of all preserved parts. Despite subsequently being treated as the specimen of reference, this fossil was not designated a neotype, however, and lacks the head, necessary for assessing family and suborder affinities (see below). Therefore, to bring clarity and nomenclatural stability to the Furagrion concept, we will be requesting in a forthcoming Case to the International Commission on Zoological Nomenclature that they designate as neotype of Phenacolestes jutlandicus specimen MM-10752, which is a well-preserved specimen in dorsal aspect, with all four wings and body almost complete, including the faint but distinct left compound eye (the right is indistinct), and parts of three legs (Figs. 3–4). *not included in PCA, lacking too many characters. Family assignment and the dysagrionoid grade. The wings of MM-10752 and the FUM-N 13856 wing are consistent with all nine wing character states used in part for the Dysagrionidae diagnosis (Archibald et al. 2021), and Henriksen’s MGUH 1819 possesses those character states for the characters that are observable. These are (see Fig. 5, shown on the Dysagrion lakesii Scudder wing): 1- crossvein O absent; 2- arculus at or closely proximad Ax2; 3- quadrangle broad, distal side longer than proximal, posterior side longer than anterior, distal-posterior angle oblique, proximal-anterior angle usually about 90°; 4- nodus at least a quarter wing length, usually more; 5- AA, AP branch before joining CuP, AA briefly free distad petiole; 6- RP3-4 originates ca. one to usually two thirds the length from arculus to subnodus; 7- antesubnodal space without crossveins (note: Nel & Jouault (2022) mistakenly read this as antenodal space); 8- CuA–A space expanded in middle, at least two cells wide; 9- CuA long, ends on posterior margin at mid-wing or further These character states define the Dysagrionidae in combination with head character states of the suborder Cephalozygoptera (diagnosis of Archibald et al. 2021): width across eyes about twice the length from the anterior margin of antefrons to the posterior of the occiput; compound eyes more or less adpressed to head capsule, convex laterally but not hemispherical, their posterolateral corners extended posteriorly to varying degrees, sometimes even acutely; the distance between compound eyes at the level of the centre of the ocelli is about the width of one eye or less, i.e., the head is not shortened and distinctly extended laterally with bulging, hemispherical compound eyes as in Zygoptera. Archibald et al. missed that Rust (1999) had found that although the compound eyes are indistinctly preserved in Furagrion specimen 16-B3618, they are present and widely separated as in Zygoptera (“Von den grossen Komplexaugen sind nur undeutliche Reste überliefert. Sie liegen, wie für Zygopteren charakteristisch, weit voneinander getrennt an den Aussenseiten des Kopfes”, p. 19). He did not illustrate this specimen, and its whereabouts is not known to us. We examined several specimens that conform to Rust’s observation and, therefore, treat the genus as a zygopteran. The compound eyes are clearly preserved in 14M-A2163 (Fig. 6) and are faintly but confidently preserved in the proposed neotype MM-10752 (Fig. 3). These have the typical zygopteran shape, hemispherical, widely set and bulging outward, and the head is short. The compound eyes are not preserved in MM-10750 (Figs. 7–8), but the remaining head capsule is short and wide as in Zygoptera. Such a loss of eyes in fossils may happen, especially in Zygoptera, as they protrude and are more fragile, apparently more easily degraded than the robust head capsule, or they might simply break away from the head in death. Such missing compound eyes can also be clearly seen in multiple specimens of Lestes ceresti Nel & Papazian from the Oligocene of Céreste, France, cf. the holotype MNHN. F.R07445 (Archibald & Cannings 2021 fig. 1B and 1C) and PNRL 2019 and PNRL 2021 (Nel & Jouault 2022, figs 11A and 12; the head of L. ceresti PNRL 2020 in their fig. 11B appears too poorly preserved to confidently evaluate) and see Chalcolestes tibetensis Xia et al. (Xia et al. 2022, figs. 3A and 4) and Nel & Zheng (2021 fig. 2B). Some other Zygoptera possess wing venation with many of the nine diagnostic character states of Dysagrionidae. For example, the extant Argia funcki Sélys (Coenagrionidae, Coenagrionoidea) only differs by character state 6, and species of Austroargiolestes Kennedy (Argiolestidae Fraser, ‘Calopterygoidea’) by 5 and 6 (both Fig. 5). The wing of the Eocene Viridiflumineagrion aasei Nel (Fig. 5) differs by character state 6 (assigned to “‘Megapodagrionidae’ sensu lato ”, which is highly polyphyletic, see Dijkstra et a. (2014), and so we treat it as family indet.). The extinct zygopteran Oligolestes Nel & Escuillé (family indet., see below) bears all character states but 1, 6, and 9 and Eodysagrion Rust et al. (Rust et al. 2008, Fig. 9) (provisionally in the Thaumatoneuridae: Huang et al. 2017) differs only by character state 3. Miopodagrion possesses a zygopteran head, but much of its overlapping and variably preserved wings are difficult to separate and interpret with confidence (Fig. 9). These wings can be established to possess character states 2, 3, 4, 8 and 9, but 1, 5, 6, and 7 appear uncertain or unknowable in its single fossil. We also consider it to be of unknown family. Treintamilun Petrulevičius (Frenguelliidae) (Fig. 5) shares all nine character states except character state 1 (absence of crossvein O), which cannot be assessed by preservation; however, Petrulevičius & Nel (2003) report this crossvein in the other described frenguelliid genus, Frenguellia Petrulevičius & Nel. Frenguelliids bear a distinctive CuP, indicating that that family might not belong to either the Zygoptera or Cephalozygoptera, but could belong to an undescribed suborder (Petrulevičius & Nel 2003, Petrulevičius 2017). Combinations of dysagrionid wing character states are then found widely across even distantly related odonates. Genera belonging to the Dysagrionidae sensu Archibald et al. (2021) or possibly so were previously assigned to a variety of extant zygopteran families by wing venation, highlighting the generalisation of these character states. These include the “calopterygoid” families Thaumatoneuridae, Pseudolestidae, Megapodagrionidae (as then defined), Amphipterygidae, and “ Agrionidae ” (= Calopterygidae) (e.g., Scudder 1878, Campion 1913, Tillyard & Fraser 1939, Fraser 1957, Carpenter 1992, Nel & Paicheler 1994, Bechly 1996, Rust 1999, Nel et al. 2005 a, 2005b; Nel & Arillo 2006, Rust et al. 2008, Garrouste & Nel 2015, Nel et al. 2016; Zheng et al. 2016a, 2016b, 2017, Huang et al. 2017). There is thus a “dysagrionoid grade” of wing venation found among Zygoptera and Cephalozygoptera, and following Petrulevičius & Nel (2003) and Petrulevičius (2017) on the status of Frenguelliidae, then even outside of these suborders. These dysagrionoid wing character states might be symplesiomorphies shared by common ancestors which date back at least to the early Jurassic (Kohli et al. 2021, Suvorov et al. 2022) with their stem taxa possessing wings that might look very much like those of e.g., the Frenguelliidae Petrulevičius & Nel (or Congqingia Zhang?). The family-level phylogeny of extant Zygoptera has undergone considerable development since the largely unresolved cladogram of Dijkstra et al. (2014) (see Bybee et al. 2021, Kohli et al. 2021, and Suvorov et al. 2022) and the superfamily Lestioidea Calvert appears to be well supported as sister group to the remaining extant Zygoptera. Within the latter, Platystictidae Kennedy are sister to the remaining non-lestoid Zygoptera (Bybee et al. 2021, Kohli et al. 2021, Suvorov et al. 2022). Neither Lestoidea nor Platystictidae include taxa that possess the dysagrionoid wing venation (e.g., Garrison et al. 2010). The presence of dysagrionoid character states in zygopteran taxa might also represent homoplastic reversals as adaptations to similar selection pressures. A thorough study of the deep-time evolution of these wing characters based on basal zygopteran-cephalozygopteran phylogeny falls well outside the scope of this paper, but should be the focus of a future study. The family and suborder designations of dysagrionoid grade taxa. The heads of Dysagrion Scudder, Phenacolestes, Petrolestes Cockerell, Congqingia, Okanagrion Archibald & Cannings, and Okanopteryx Archibald & Cannings, are known and consistent with the Dysagrionidae and Cephalozygoptera concepts (Archibald et al. 2021; Archibald & Cannings 2021) and so are confidently established in that family and suborder. Archibald et al. (2021) further mention the presence of antenodal crossvein Ax0 as a potentially important character for identifying possible Cephalozygoptera fossils where the head is unknown. Although the wing base where Ax0 is found is often poorly preserved or absent in fossils, the vein is found in Dysagrionidae and Sieblosiidae where this region is well-preserved, and also found in the Whetwhetaksidae Archibald & Cannings, strengthening the notion that they are closely related (Archibald et al. 2021, Simonsen et al. 2022). Ax0 is absent or covered by sclerotization in Zygoptera (Bechly 1996, Rehn 2003), except found in two Eocene species of Euphaeidae Jacobson & Bianchi (Archibald et al. 2012) and Burmadysagrion zhangi Zheng et al. (Zheng et al. 2016a). The wing base is well preserved in several of the fossils studied here (e.g., Figs 2, 16-18, 21), and although we find sclerotization in the region (e.g., Fig. 21), we do not find evidence for the presence of Ax0. The family affinities of Furagrion and Viridiflumineagrion are unknown, as their dysagrionoid wing venation (poorly known in Miopodagrion) alone is insufficient to establish this, and other relevant characters are little known beyond aspects of their heads that establish them as zygopterans. The family affinity of Oligolestes Schmidt is also unknown (Nel et al. 2005a; Nel et al. 2020; Nel & Zheng 2021; Nel & Jouault 2022). These authors compared it to the Sieblosiidae Handlirsch, but excluded that family as its diagnosis (Nel et al. 2005a, p. 223) consists of a single character state that Oligolestes lacks: “highly specialised nodus apparently traversed by ScP, as the terminal kink of CP is shifted basally together with the nodal and subnodal veinlets and the nodal membrane sclerotisation is reduced”. Nel et al. (2005a, p. 223) concluded that “ Oligolestes could be closely related to the Sieblosiidae sensu stricto, but there is no proof supporting this hypothesis because all these characters are individually present in other damselfly lineages.” We agree with Nel & Jouault (2022) that Oligolestes stoeffelensis Nel et al. is a zygopteran by its distinctly zygopteran head and with all of the above authors that the genus cannot be assigned to a family given current knowledge. Further, the Cephalozygopteran head is clearly seen in the Sieblosiidae (see Stenolestes Scudder: Stenolestes cf. fischeri Nel MNHNF-B.47288 and less clearly in Stenolestes falloti (Théobald) holotype MNHN.F.B24507, Archibald & Cannings 2021, fig. 2a, b). Odonates with dysagrionoid wings where the head is unknown are also then family and suborder indet., and at most might be considered cf. Cephalozygoptera, Dysagrionidae. These include Primorilestes Nel et al.; Electrophenacolestes Nel & Arillo; Stenodiafanus Archibald & Cannings; Menatagrion Nel & Jouault; Allenbya Archibald & Cannings; Thanetophilosina Nel et al.; Valerea Garrouste et al.; the unnamed Alaskan Chickaloon specimen of Garrouste & Nel (2019); specimen MeI6572 of the Senckenberg Museum, Frankfurt, Germany, treated as Megapodagrionidae genus and species A by Petrulevičius et al. (2008) and as cf. Dysagrionidae gen and sp. A by Archibald et al. (2021); specimen NHMUK I.9866/I.9718 of Nel & Fleck (2014). The Whetwhetaksidae is cf. Cephalozygoptera with more confidence by the presence of Ax0 (see above). Principle component analysis and variation within the genus. PCA results are shown in Fig. 10. The “ratios” plot (Fig. 10A) is based on the eight calculated ratios only, i.e., is based on wing sh, Published as part of Archibald, S. Bruce, Ware, Jessica L., Rasmussen, Jan A., Sylvestersen, René L., Olsen, Kent & Simonsen, Thomas J., 2023, The damselfly genus Furagrion Petrulevičius et al. (Odonata, Zygoptera) from the early Eocene Fur Formation of Denmark and the dysagrionoid grade, pp. 289-317 in Zootaxa 5278 (2) on pages 292-302, DOI: 10.11646/zootaxa.5278.2.4, http://zenodo.org/record/7906172, {"references":["Zessin, W. (2011) Neue Insekten aus dem Moler (Palaozan / Eozan) von Danemark Teil 1 (Odonata: Epallagidae, Megapodagrioniidae), Virgo, Mitteilungsblatt des Entomologischen Vereins Mecklenburg, 14, 64 - 73.","Henriksen, K. (1922) Eocene insects from Denmark. Danmarks Grologiske UndersOgelser, II Raekke, 37, 1 - 36.","Storey, M., Duncan, R. A. & Swisher, C. C. (2007) Paleocene-Eocene Thermal Maximum and the Opening of the Northeast Atlantic. Science, 316 (5824), 587 - 589. https: // doi. org / 10.1126 / science. 1135274","Stokke, E. W., Jones, M. T., Tierney, J. E., Svensen, H. H. & Whiteside, J. H. (2020) Temperature changes across the Paleocene-Eocene Thermal Maximum - a new high-resolution TEX 86 temperature record from the Eastern North Sea Basin. Earth and Planetary Science Letters, 544, 116388. https: // doi. org / 10.1016 / j. epsl. 2020.116388","Nel, A. & Jouault, C. (2022) The odonatan insects from the Paleocene of Menat, central France. Acta Palaeontologica Polonica, 67, 631 - 648. https: // doi. org / 10.4202 / app. 00960.2021","Garrouste, R. & Nel, A. (2019) Alaskan Palaeogene insects: a challenge for a better knowledge of the Beringian ' route' (Odonata: Aeshnidae, Dysagrionidae). Journal of Systematic Palaeontology, 17, 1939 - 1946. https: // doi. org / 10.1080 / 14772019.2019.1572235","Petrulevicius, J. F., Wappler, T., Wedmann, S., Rust, J. & Nel, A. (2008) New Megapodagrionid Damselflies (Odonata: Zygoptera) from the Paleogene of Europe. Journal of Paleontology, 82, 1173 - 1181.","Cockerell, T. D. A. (1908 a) Fossil insects from Florissant, Colorado. Bulletin of the American Museum of Natural History, 24, 59 - 69.","Nel, A. & Paicheler, J. - C. (1994) Les Lestoidea fossiles. Un inventaire critique (Odonata, Zygoptera). Annales de Paleontologie, 90, 1 - 59. (rang B)","Rust, J. (1999) Biologie der Insekten aus dem altesten Tertiar Nordeuropas. Habilitationsschrift zur Erlangung der venia legendi fu ¨ r das Fach Zoologie. Postdoctoral thesis, Biologische Fakultat der Georg-August-Universitat Gottingen, Gottingen, Lower Saxony, 482 pp.","Xia, G., Zheng, D., Krieg-Jacquier, R., Fan, Q., Chen, Y. & Nel, A. (2022) The oldest-known Lestidae (Odonata) from the late Eocene of Tibet: palaeoclimatic implications. Geological Magazine, 159, 511 - 518. https: // doi. org / 10.1017 / S 0016756821001102","Nel, A. & Zheng, D. (2021) The recently proposed odonatan ' suborder' Cephalozygoptera: fact or fiction. Palaeoentomology, 004, 165 - 170. https: // doi. org / 10.11646 / palaeoentomology. 4.2.5","Rust, J, Petrulevicius, J. F. & Nel, A. (2008) The first damselflies from the lowermost Eocene of Denmark, with a description of a new subfamily (Odonata, Zygoptera, Dysagrionidae). Palaeontology, 51, 709 - 713. https: // doi. org / 10.1111 / j. 1475 - 4983.2008.00780. x","Huang, D., Azar, D., Cai, C., Maksoud S., Nel, A. & Bechly, G. (2017) Mesomegaloprepidae, a remarkable new damselfly family (Odonata: Zygoptera) from mid-Cretaceous Burmese amber. Cretaceous Research, 73, 1 - 13. https: // doi. org / 10.1016 / j. cretres. 2017.01.003","Petrulevicius, J. F. & Nel, A. (2003) Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata. Journal of Natural History, 37 (24), 2909 - 2917. https: // doi. org / 10.1080 / 0022293021000007543","Petrulevicius, J. F. (2017) First Frenguelliidae (Insecta: Odonata) from the middle Eocene of Rio Pichileufu, Patagonia, Argentina. Arquivos Entomoloxicos, 18, 367 - 374.","Scudder, S. H. (1878) An account of some insects of unusual interest from the Tertiary rocks of Colorado and Wyoming. Bulletin of the United States Geological and Geographical Survey of the Territories, 4, 519 - 543.","Campion, H. (1913) The antenodal reticulation of the wings of agrionine Dragonflies. Proceedings of the Academy of Natural Sciences of Philadelphia, 65, 220 - 224.","Tillyard, R. J. & Fraser, F. C. (1939) A reclassification of the order Odonata based on some new interpretations of the dragonfly wing by R. J. Tillyard, continuation thereof by F. C. Fraser, Part II, the suborder Zygoptera (continued), Protanisoptera. Australian Zoologist, 9, 195 - 221.","Fraser, F. C. (1957) A reclassification of the order Odonata, Based on Some New Interpretations of the Venation of the Dragonfly Wing By the late R. J. Tillyard and F. C. Fraser, Revision. Royal Zoological Society of New South Wales, Sydney, 133 pp.","Carpenter, F. M. (1992) Superclass Hexapoda. In: Kaesler, R. L. (Ed.), Treatise on Invertebrate Paleontology. Part R. Arthropoda 4. Vols. 3 & 4. The Geological Society of America, Boulder, Colorado & University of Kansas, Lawrence, Kansas, pp. 1 - 655.","Bechly, G. (1996) Morphologische Untersuchungen am Flu ¨ gelgeader der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata) unter besonderer Beru ¨ chsichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Boblingen, Special Volume 2, 1 - 402.","Nel, A., Petrulevicius, J. F., Gentilini, G. & Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblosiidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007","Nel, A., Petrulevicius J. F. & Jarzembowsk, E. A. (2005 b) New fossil Odonata from the European Cenozoic (Insecta: Odonata: Thaumatoneuridae, Aeshnidae,? Idionychidae, Libellulidae). Neues Jahrbuch fu ¨ r Geologie und Palaontologie, Abhandlungen, 235, 343 - 380.","Nel, A. & Arillo, A. (2006) The first Baltic amber dysagrionine damselfly (Odonata: Zygoptera: Thaumatoneuridae: Dysagrioninae). Annales de la Societe entomologique de France, New Series, International Journal of Entomology, 42, 179 - 182. https: // doi. org / 10.1080 / 00379271.2006.10700621","Garrouste, R. & Nel, A. (2015) New Eocene damselflies and the first Cenozoic damseldragonfly of the isophlebiopteran lineage (Insecta: Odonata). Zootaxa, 4028 (3), 354 - 366.","Nel, A., Simov, N., Bozukov, V. & Marinov, M. (2016) New dragonflies and damselflies from Middle Miocene deposits in SW Bulgaria (Insecta: Odonata). Palaeontologia Electronica, 19.3.35 A, 1 - 13. https: // doi. org / 10.26879 / 642","Zheng, D., Wang, B. Jarzembowski, E. A. Chang, S. - C. & Nel, A. (2016 a) Burmadysagrioninae, a new subfamily (Odonata: Zygoptera: Dysagrionidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 67, 126 - 132. https: // doi. org / 10.1016 / j. cretres. 2016.07.006","Zheng, D., Zhang, Q., Nel, A., Jarzembowski, E. A., Zhou, Z., Chang, S. - C. & Wang, B. (2016 b) New damselflies (Odonata: Zygoptera: Hemiphlebiidae, Dysagrionidae) from mid-Cretaceous Burmese amber. Alcheringa, 41, 12 - 21. https: // doi. org / 10.1080 / 03115518.2016.1164402","Zheng, D., Chang, S. - C., Nel, A., Jarzembowski, E. A., Zhuo, D. & Wang, B. (2017) Electrodysagrion lini gen. et sp. nov., the oldest Dysagrionini (Odonata: Zygoptera: Dysagrionidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 77, 44 - 48. https: // doi. org / 10.1016 / j. cretres. 2017.05.008","Tann, J. (2021) Images of Australian Odonata Wings. Technical Reports of the Australian Museum Online, 33, 1 - 101. https: // doi. org / 10.3853 / j. 1835 - 4211.33.2021.1767","Garrison, R. W., von Ellenrieder, N. & Louton, J. A. (2010) Damselfly genera of the New World: an illustrated and annotated key to the Zygoptera. Johns Hopkins University Press, Baltimore, Maryland, 490 pp.","Kohli, M., Letsch, H., Greve, C., Bethoux, O., Deregnaucourt, I., Liu, S., Zhou, X., Donath, A., Mayer, C., Podsiadlowski, L., Gunkel, S., Machida, R., Niehuis, O., Rust, J., Wappler, T., Yu, X., Misof, B. & Ware, J. (2021) Evolutionary history and divergence times of Odonata (dragonflies and damselflies) revealed through transcriptomics. iScience, 24, 103324.","Suvorov, A., Scornavacca, C., Fujimoto, M. S., Bodily, P., Clement, M., Crandall, K. A., Whiting, M. F., Schrider, D. R. & Bybee, S. M. (2022) Deep ancestral introgression shapes evolutionary history of dragonflies and damselflies. Systematic Biology, 71, 526 - 546. https: // doi. org / 10.1093 / sysbio / syab 063","Dijkstra, K. - D. B., Kalkman, V. J., Dow, R. A., Stokvis, F. R. & van Tol, J. (2014) Redefining the damselfly families: the first comprehensive molecular phylogeny of Zygoptera (Odonata). Systematic Entomology, 39, 68 - 96. https: // doi. org / 10.1111 / syen. 12035","Bybee, S. M., Kalkman, V. J., Erickson, R. J., Frandsen, P. B., Breinholt, J. W., Suvorov, A., ... & Ware, J. L. (2021). Phylogeny and classification of Odonata using targeted genomics. Molecular phylogenetics and evolution, 160, 107115. https: // doi. org / 10.1016 / j. ympev. 2021.107115","Simonsen, T. J., Archibald, S. B., Rasmussen, J. A., Sylvestersen, R. L., Olsen, K. & Ware, J. L. (2022) Danowhetaksa gen. Nov. With two species from the early Eocene Olst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera). Zootaxa, 5099 (5), 586 - 592. https: // doi. org / 10.11646 / zootaxa. 5099.5.5","Rehn, A. C. (2003) Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology, 28, 181 - 239. https: // doi. org / 10.1046 / j. 1365 - 3113.2003.00210. x","Nel, A., Poschmann, M. J. & Wedmann, S. (2020) New dragonflies and damselflies (Odonata) from the late Oligocene of Enspel (Rhineland-Palatinate, SW Germany). Palaeontologia Electronica, 23 (3), a 59. https: // doi. org / 10.26879 / 1126","Nel, A. & Fleck, G. (2014) Dragonflies and damselflies (Insecta: Odonata) from the Late Eocene of the Isle of Wight. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 104, 283 - 306. https: // doi. org / 10.1017 / S 175569101400005 X"]}

Floricimbex vetusculus (Cockerell, 1922) Fig. 27. Cimbex vetusculus Cockerell, 1922, p. 50. Holotype. DMNH-163A, B, type specimen, Cockerell (1922). Part, a rather complete specimen; counterpart, missing the right wings and head. Diagnosis. As for genus. Description. Holotype, male. Body: length 16.7 mm as preserved, colour pattern possibly mostly dark (organic matter mostly lost, obscuring colouration). No rough surface sculpture discernible. Structure of head and thorax obscure. Mid, hind coxae touching basally, slightly separated distally, hind coxae large, elongate, more or less parallel-sided, no distinct armament visible; hind femur almost regularly elliptical, 4.4 mm long, 1.9 mm wide (length to abdominal segment width ca. 1:2.3), other leg segments incomplete. Wings damaged, incompletely known. Forewing length 13.9 mm, maximum width 4.4 mm or slightly more, length/width> 3.1; membrane possibly lightly infuscate behind R+M; pterostigma dark; R+M shorter than 1-M; maximum length of cell 2r shorter than 3r; 2r-rs slightly longer than Rs between 2r-m, 2r-rs, joining cell 3rm slightly before its middle; 2r-m straight, joins Rs distal to 2m-cu joining M, well distal 2mcu; 1-M almost straight, sub-parallel to 1m-cu; 3r-m incompletely preserved, its form obscure; cell 2mcu with M apparently subequal to 1m-cu; 2m-cu incompletely preserved, apparently slightly curved toward base; cu-a interstitial; anal cells not preserved. Hind wing with M base basal to level of Rs base, m-cu present, cu-a before middle of cell mcu, a1-a2 basal to level of Rs base, Cu with distinct free apex, 1A and 2A with long free apex. Abdomen narrow, slightly widening caudally (widest at segments 5–6), tergum 1 with straight hind margin, genitalia not distinct. Material. Holotype. Locality and age. Florissant Formation, Priabonian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 30-31, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Cockerell, T. D. A. (1922) The fossil sawflies of Florissant, Colorado. The Entomologist, 55, 49 - 50."]}

Genus Pseudocimbex Rohwer, 1908 Fig. 28. Pseudocimbex Rohwer, 1908, p. 526. Type species: Pseudocimbex clavatus Rohwer, 1908, by original designation. Diagnosis. Distinct from all other Symphyta by hind wing with M+Cu branching to M, Cu unusually basally; M and Cu at a low angle to M+Cu, almost aligned with it; cell mcu long, narrow, wedge-shaped; cu-a present, vertical. Also distinct from all other Symphyta by comparatively short antenna (about as long as half head width) with subspherical club consisting of several very short segments. Additionally distinct from Cimbicidae by lack of 2r-rs; by its wide costal space; and by presence of vein Sc. Remarks. 2r-rs is not detected in the forewing in some cimbicid specimens described here, presumably by preservation. It is, however, confidently absent in both forewings of Pseudocimbex examined dry and wetted with ethanol (Fig. 28). The very short antenna does occur in various Pergidae, however, there it is combined with an elongate and non-segmented club. In comparison with extant species, a placement of this fossil in either Cimbicidae or Pergidae remains ambiguous. Species included. Type species only., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 31-32, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Rohwer, S. A. (1908) On the Tenthredinoidea of the Florissant Shales. Bulletin of the American Museum of Natural History, 24, 521 - 530."]}

Allenbycimbex n. gen. http://zoobank.org/ urn:lsid:zoobank.org:act: 704AB0FB-18D0-4CF4-AB78-F48D42378EAF Fig. 4. Type species. Allenbycimbex morrisae n. sp. Diagnosis. Distinguished from Cenocimbex by R lacking dark triangular sclerite before base of pterostigma reaching 1-Rs+M near 1m-cu; wing narrower (length/width ca. 3.0 as reconstructed) [Cenocimbex: 2.5]; cell 3rm long (2rm to 3r-m distinctly longer than length of 3r-m); hind wing with m-cu placed much basally than in Cenocimbex; distinguished from Leptostigma by pterostigma broad, maximum height distinctly exceeding height of C to R basal to pterostigma. Locality and age. Vermilion Bluffs Member of the Allenby Formation south of the village of Princeton, British Columbia, Canada; mid-Ypresian. Species included. Type only. Etymology. Named for the Allenby Formation, where its only specimen was found. The name is to be treated with masculine grammatical gender since it is derived from Cimbex (see Taeger et al. 2010: 188).

Trichiosomites obliviosus Brues, 1908 Figs. 24–25. Trichiosomites obliviosus Brues, 1908 p. 260, Fig. 1. Holotype. Holotype MCZ 2036, Brues (1908). A rather complete female with head present, but positioned under thorax (not missing as Brues 1908 presumed). Diagnosis. As for genus. Description. Holotype MCZ 2036 (Fig. 24), female. Body: head length 0.5 mm or slightly more, width 0.87 mm; thorax length ca. 2.5 mm; abdomen length ca. 5 mm; body colouration unknown by preservation. Forewing: length ca. 6.0 mm, estimated maximum width ca. 2.0 mm, length/width: ca. 3.0. UCM 4570 (Fig. 25), female. Body: length ca. 9.2 mm; head, antenna dark; thorax dark, wings including pterostigma light; abdomen light. Head difficult to interpret as preserved; moderately transverse, occiput long, three distinct ocelli in low triangle, compound eyes not preserved, sole funicular segment thin, slightly more than half length of club; club expands gradually over three segments (“drop-like”). Thoracic dorsum only partially visible as preserved, median longitudinal mesonotal sulcus and notauli present; mesopleuron with no mesopleural sulcus visible. Forewing: membrane hyaline, length ca. 6.5 mm; maximum width estimated ca. 2.5 mm, length/width: ca. 2.6; pterostigma wide, ca. as cell 2r; R+M shorter than 1-M; maximum length of cell 2r shorter than 3r; 2r-rs longer than Rs between 2r-m, 2r-rs, joining Rs near middle of cell 3rm; 2r-rm straight, short, joins M at 2m-cu; 1-M curved; cell 1mcu with M, 1m-cu sub-parallel; 3r-m straight; cell 2mcu with M slightly longer than 1m-cu; 2m-cu straight; cu-a postfurcal; anal cell separated by 1A fusion with 2A+3A (no crossvein) for length of about a third length of cell 2a. Abdomen: long, wide; ovipositor thick, not exceeding abdomen apex. Material. Holotype and UCM 4570, a rather complete specimen missing only the legs and much of the hind wings. Locality and age. Florissant Formation, Priabonian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 27-28, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Brues, C. T. (1908) New Phytophagous Hymenoptera from the Tertiary of Florissant, Colorado. Bulletin of the Museum of Comparative Zoology, Harvard College, 51, 259 - 276. https: // doi. org / 10.5962 / bhl. title. 19923"]}

Genus Cenocimbex Nel, 2004 Figs. 2, 3E–F. Type species C. menatensis Nel, 2004. Emended diagnosis. Separated from all other Cenocimbicinae by R before base of pterostigma with dark triangular sclerite reaching 1-Rs+M near 1m-cu [vs. sclerotization not apparent, either absent or pale, small]; by wing broad, estimated length/width 2.5 [all others ca. 2.8 or more]; by cell 3rm short (2r-m to 3r-m length ≤ 3r-m [vs. distinctly longer]); from Leptostigma n. gen. also by pterostigma broad, maximum height distinctly more than height of C to R basal to pterostigma [Leptostigma n. gen.: similar in height to C to R basal to pterostigma]. Hind wing with r-m, m-cu and cu-a only moderately distant from each other (for about length of m-cu, so cu-a at about midlength cell mcu) [in other genera usually well before mid-length]. Remarks. Sclerotisation on R near the pterostigma, even though of much lesser size, is clearly seen in an extant cimbicine, see Vilhelmsen et al. 2018, Fig. 2B). Locality and age. Menat Formation; Puy-de-Dôme, France; Selandian. Species included. Type species only.

Cenocimbicinae and the history of Cimbicidae The Cimbicidae first appears in the fossil record with one fossil of the extinct subfamily Cenocimbicinae in the Selandian, which is dominant and diverse in the Ypresian and disappears after. A single specimen of the extant Pachylostictinae is known in the Ypresian. A small assemblage of the extant subfamilies Abiinae and Cimbicinae appears in the late Priabonian, the family is unknown the Oligocene, and then it reappears in the Miocene showing the modern pattern of dominance by the Cimbicinae and Abiinae (Table 1). The family might have originated in the Paleocene, but this might also simply reflect the paucity of Paleocene and Late Cretaceous shale insect localities. They—and Symphyta in general—are almost entirely absent in amber for an unknown reason. Their rich record in the Okanagan Highlands could be an artefact of unknown taphonomic factors favouring their preservation there. ......continued on the next page *Misspelled chromptera by Taeger et al. (2010, #2263) Alternatively, this record might closely reflect their actual history (see Archibald et al. 2018). Cimbicidae feed on the leaves of angiosperms: Abiinae and Cimbicinae on Betulaceae, Caprifoliaceae, Dipsacaceae, Rosaceae, and Salicaceae; and the Corynidinae on Crassulaceae, Geraniaceae and Papaveraceae (the hosts of Pachylostictinae are little known) (summarized by Vilhelmsen 2019, see references therein). All of these are present in the Okanagan Highlands except the Dipsacaceae and Geraniaceae, and many genera of these families that are important today first appear there (e.g., Crane & Stockey 1987, Wolfe & Wehr 1987; Wehr & Hopkins 1994; Greenwood et al. 2005; Moss et al. 2005; Devore & Pigg 2007, 2010; Mathewes et al. 2016; Pigg & Devore 2016). The temperate montane Okanagan Highlands had a high diversity of insects and plants (woody dicots analysed) associated with climatic factors (Archibald et al. 2010), and may have been the setting for the origin of modern microthermal northern forests that the great majority of extant Cimbicidae prefer which later spread downslope into lowlands in the globally cooler Neogene (Graham 2011). The only fossil Pachylostictinae known is from the Ypresian megathermal lowland Green River Formation. The Priabonian record of the Cimbicidae (excluding the single Baltic amber larva) is at Florissant, another upper microthermal montane forest, where only modern subfamilies are present. Such montane forest communities are rare in the insect fossil record, which might further bias our knowledge of the history of the family. The limited morphological information available in known fossils indicates that the Cenocimbicinae forms a clade with the Cimbicinae and Abiinae based on at least two synapomorphies of the forewing: 1, cell 2r together with the pterostigma is distinctly higher than cell 3r (Corynidinae and Pachylostictinae: at most as high as 3r), and 2, crossvein 2r-m joins M distal of 2m-cu (basal in Corynidinae and Pachylostictinae). The strongly bent crossvein 2r-m of Cenocimbicinae separates it from the Cimbicinae + Abiinae, where it is straight or gently curved (we treat rare occurrences of a similar 2r-m in Cimbicinae and Abiinae as convergent). This implies an unknown presence of Corynidinae and Pachylostictinae at least to the Selandian, and of Cimbicidae originating earlier in the Paleocene or perhaps the Late Cretaceous., Published as part of Archibald, S. 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EPHE Biologie et Evolution des Insectes, Paris, 5, 115 - 120, 4 fig.","Steinbach, G. (1967) Zur Hymenopterenfauna des Pliozans von Willershausen / Westharz. Bericht der Naturhistorischen Gesellschaft zu Hannover, 111, 95 - 102.","Archibald S. B., Rasnitsyn, A. P., Brothers, D. J. & Mathewes, R. W. (2018) Modernisation of the Hymenoptera: ants, bees, wasps and sawflies of the early Eocene Okanagan Highlands. The Canadian Entomologist, 150, 205 - 257. https: // doi. org / 10.4039 / tce. 2017.59","Vilhelmsen, L. (2019) Giant sawflies and their kin: morphological phylogeny of Cimbicidae (Hymenoptera). Systematic Entomology, 44, 103 - 127. https: // doi. org / 10.1111 / syen. 12314","Crane, P. R. & Stockey, R. A. (1987) Betula leaves and reproductive structures from the Middle Eocene of British Columbia, Canada. Canadian Journal of Botany, 65, 2490 - 2500. https: // doi. org / 10.1139 / b 87 - 338","Wolfe, J. A. & Wehr, W. C. (1987) Middle Eocene dicotyledonous plants from Republic, northeastern Washington. US Geological Survey Bulletin 1597, 1 - 25.","Wehr, W. C. & Hopkins D. Q. (1994) The Eocene orchards and gardens of Republic, Washington. Washington Geology, 22, 27 - 34.","Greenwood, D. R., Archibald, S. B., Mathewes, R. W. & Moss, P. T. (2005) Fossil biotas from the Okanagan Highlands, southern British Columbia and northern Washington State: climates and ecosystems across an Eocene landscape. Canadian Journal of Earth Sciences, 42, 167 - 185. https: // doi. org / 10.1139 / e 04 - 100","Moss, P. T., Greenwood, D. R. & Archibald, S. B. (2005) Regional and local vegetation community dynamics of the Eocene Okanagan Highlands (British Columbia-Washington State) from palynology. Canadian Journal of Earth Sciences, 42, 187 - 204. https: // doi. org / 10.1139 / e 04 - 095","DeVore, M. L. & Pigg, K. B. (2007) A brief review of the fossil history of the family Rosaceae with a focus on the Eocene Okanogan Highlands of eastern Washington State, USA, and British Columbia, Canada. Plant Systematics and Evolution, 266, 45 - 57. https: // doi. org / 10.1007 / s 00606 - 007 - 0540 - 3","DeVore, M. L. & Pigg, K. B. (2010) Floristic composition and comparison of middle Eocene to late Oligocene floras of North America. Bulletin of Geosciences, 85, 111 - 134. https: // doi. org / 10.3140 / bull. geosci. 1135","Mathewes, R. W., Greenwood, D. G. & Archibald, S. B. (2016) Paleoevironment of the Quilchena Flora, British Columbia, during the Early Eocene climatic optimum. Canadian Journal of Earth Sciences, 53, 574 - 590. https: // doi. org / 10.1139 / cjes- 2015 - 0163","Pigg, K. B. & Devore, M. L. (2016) A review of the plants of the Princeton chert (Eocene, British Columbia, Canada). Botany, 94, 661 - 681. https: // doi. org / 10.1139 / cjb- 2016 - 0079","Graham, A. (2011) The age and diversification of terrestrial New World ecosystems through Cretaceous and Cenozoic time. American Journal of Botany, 98, 336 - 351. https: // doi. org / 10.3732 / ajb. 1000353"]}

Genus Eopachylosticta Malaise, 1945 Eopachylosticta Malaise, 1945, p. 14. Type species. Amasis byrami Cockerell, 1925. Emended diagnosis. Because of lack of sufficiently preserved wings and incompletely preserved body, the type and only fossil lacks many characters reliably used as diagnostic in Cimbicidae. However, generic distinction in Pachylostictinae is currently based in part on colour pattern (Vilhelmsen et al. 2019) which is available here, allowing at least a tentative genus diagnosis until better preserved fossils are found. The genus is distinct from others of Pachylostictinae by its dark head and thorax in contrast to its distinctly paler (somewhat infuscate) abdomen. Remarks. Eopachylosticta belongs to the Pachylostictinae by a combination of its antennae with five segments and the median longitudinal mesonotal sulcus and notauli present and well developed (Vilhelmsen 2019). Preservation of the type specimen of Eopachylosticta byrami suffers from the aggressive hydrochemistry of the Green River depositional environment. This is a common problem for fossil insects in parts of this formation, where the wings, legs and other body parts which were pale in life may become extremely faint or invisible as fossils. Therefore, our description and interpretation of this fossil is limited to distinguishing the genus from others of Pachylostictinae, setting aside more detailed analysis until better preserved material is available. Species included. Type species only., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 24-25, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Malaise, R. (1945) Tenthredinoidoa of south-eastern Asia with a general zoogeographical review. Opuscula Entomologica Suppl, 4 (288), 1 - 288. https: // doi. org / 10.1080 / 11035894509446460","Cockerell, T. D. A. (1925) Fossil insects in the United States National Museum. Proceedings of the United States National Museum, 64, 1 - 15. [https: // www. biodiversitylibrary. org / item / 32410 # page / 7 / mode / 1 up]","Vilhelmsen, L. (2019) Giant sawflies and their kin: morphological phylogeny of Cimbicidae (Hymenoptera). Systematic Entomology, 44, 103 - 127. https: // doi. org / 10.1111 / syen. 12314"]}

Pseudocimbex clavatus Rohwer, 1908 Fig. 28. Pseudocimbex clavatus Rohwer, 1908, p. 527, Fig. 1a, e. Holotype. UCM 18614, Rohwer (1908). An almost complete specimen missing most of the legs, one antenna. Diagnosis. As for genus. Description. Holotype, female. Body: length 17.6 mm as preserved; head, thorax darker than abdomen. Surface sculpture not discernible. Structure of head, thorax obscure. Antenna very short (flagellum 1.8 mm long, less than half head width), capitate, apparently 11-segmented, scape and pedicel very short, with funicle of four segments gradually shortening distally from twice as long as wide up to strongly transverse and almost round club of possibly five segments, four very short and apical one longer. Hind tibia 4.6 mm long, other leg segments incompletely or not preserved. Forewing length 12.9 mm, maximum width 5.5 mm, length/width 2.4; membrane lightly infuscate, more so toward base; pterostigma dark, narrow, short, wedge-shaped; C, R basal to pterostigma thin, widely separated, slightly more so distally, with crossvein-like Sc between; pterostigma small, wedge-shaped; 1-M at or very close to Rs+M on R; 1r-rs short, 2r-rs lost; Rs+M very short; 2-Rs (separating cells 1r and 2rm) possibly detected (as dotted line in Fig. 28); Rs distal of 1r smoothly curved, meeting R 1 at wing margin; 2r-rs absent; 2r-m, 3r-m present, well spaced, straight, near 0.4 and 0.7 length of cell 3r; M notably angled at 1m-cu; 1m-cu short, just more than half length of 1-M, diverging with 1-M toward anterior margin, 1m-cu and 2m-cu join cells 2rm and 3rm, respectively; 1m-cu converging to 1-M forward; 2m-cu slightly curved toward base; cu-a interstitial; anal cells incompletely preserved: cell 1a apparently open posteriorly, distally (2+3A not reaching a1-a2 crossveins), cell 2a closed basally, not preserved distally (distal portion of 2+3A obscured by fore margin of hind wing). Hind wing partially known, with unusual venation; uniformly infuscate as forewing; 1r-m about its length distant from Rs base; with Rs, M+Cu, M, Cu, cu-a and 1A detected, M+Cu forking far basally of RS base, with fork narrow, M and Cu both almost aligned with M+Cu; cu-a slightly reclival, long, placed slightly basal of Rs base; 2r-m, m-cu not evident; 1A evident, with at least short free end preserved. Abdomen moderately narrow, with few details discernible; ovipositor short (length 0.65 mm), barely exceeding abdomen apex. Material. Holotype. Locality and age. Florissant Formation, Priabonian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on page 32, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Rohwer, S. A. (1908) On the Tenthredinoidea of the Florissant Shales. Bulletin of the American Museum of Natural History, 24, 521 - 530."]}

Leptostigma longitenebricum n. sp. http://zoobank.org/ urn:lsid:zoobank.org:act: 82407841-CBF9-4647-A04B-A12A242B7AE0 Figs. 3I–J, 16–18. Holotype. F-1565 (part, counterpart) (Figs. 3I–J, 16). Almost complete, in dorsal aspect, but legs, one antenna, body mostly poorly preserved, hind wings poorly preserved; from McAbee, collector John Leahy; housed in the collections of TRU. Diagnosis. Distinguished from other species of Leptostigma by antenna with a combination of club elongate, darker than flagellar base, base two long, pale segments; forewing lightly, evenly infuscate throughout. Tentatively diagnostic: R+M longer than 1-M, 2r-rs enters cell 3rm at or before its middle, 2r-m enters M far behind 2m-cu, anal cells separated by 1A joining 2A+3A for at least half height of cell 2a. Description. Holotype. Body length ca. 15.5 mm, body, legs, pterostigma dark (tarsi less dark), antenna with club darkened, funicle paler, wing membrane lightly infuscate throughout. Head roundish, eyes occupying most of head sides, oval, converging ventrally, with temples weak (scarcely visible in front view), toruli close to each other. Antenna with eight antennomeres, club elongate, narrowed apically, two basal flagellar segments cylindrical, next three widened, apical one narrowed distally; basal, apical segments longest, penultimate shortest.Pronotum with wide lateral lobes; median longitudinal mesonotal sulcus, notauli well developed, meeting well before scarcely transverse scutellum; metanotum rather narrow, with wide triangular scutellum, small, roundish cenchri. Forewing length holotype 15.5 mm (F-1563, 11.5 mm), maximum width estimated 3.5–5 mm, length/width: 3.1–3.3; pterostigma narrow, tapering, similar to width of C-R before pterostigma; R+M longer than 1-M; maximum length of cell 2r shorter than 3r; 2r-rs shorter than Rs between 2r-m, 2r-rs, joining Rs at middle of cell 3rm; 2r-rm curved, sometimes slightly sigmoidal, joins Rs at or slightly distal to 2m-cu joins M, joins M behind 2m-cu more than for half length of 2r-m; 1-M curved or (GSC 142842) almost straight; cell 1mcu with M, 1m-cu slightly divergent toward anterior margin; 3r-m near straight or (GSC 142842) distinctly bowed toward base; cell 2mcu with M similar length or (GSC 142842) shorter than 1m-cu; 2m-cu slightly curved toward base; cu-a postfurcal (holotype) or interstitial; anal cell separated by 1A fusion with 2A+3A (no crossvein) for length of almost half width of cell 2a in GSC 142842, for more than its width in F-1563. Mid, hind femora long, oval, ca. 3.5 times as long as wide; tibiae thin, only slightly longer than their femora, basitarsi ca. 0.35 as long as their tibiae. First abdominal tergum if correctly identified (F-1563 only) entire, with straight hind margin. Remarks. Differs from L. longiclava n. sp. and L. longipallidum n. sp. by flagellar base pale and, venationally, in more the distal position of 2r-rs (behind middle of cell 3rm). Differs from L. brevilatum n. sp. by its antennal club being long, also venationally by its R+M long and anal cells distinctly separated by fusion of 1A with 2A+3A. Differs from L. proxivena n. sp., L. fasciatum n. sp., L. alaemacula n. sp. with unknown antenna by its wing membrane lightly, evenly infuscate throughout (without fascia or macula) and by 2r-m entering M far behind, not close to, 2m-cu. Material. Holotype and tentatively from venation F-1563 (Fig. 17), collector unknown, in the collections of TRU, and GSC 142842, collector SBA at McAbee (GSC locality V-016800), July 12, 2000 (Fig. 18), housed in the collections of the GSC (Vancouver). Etymology. From the Latin longus, meaning long, and tenebricus, dark, referring to the antennal club. The epithet is an adjective. Locality and age. McAbee, British Columbia, Canada, Hoodoo Face beds; mid-Ypresian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 18-20, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826

Eopachylosticta byrami (Cockerell, 1925) Fig. 23. Amasis byrami Cockerell, 1925, p. 10, plate 1, Fig. 4 Eopachylosticta byrami Malaise, 1945, p. 14. Holotype. USNM 69181. A complete female with its body mostly well preserved, but almost no legs evident and poorly preserved wings with venation too faint to interpret. Green River Formation, about 8 miles south of De Beque, Colorado (Cockerell 1925), Ypresian. Diagnosis. As for genus. Description. Body length ca. 8.2 mm; head, antennae dark; thorax dark, hind femora somewhat infuscate (otherwise legs not visibly preserved, presumably pale), anterior abdomen light, posterior segments lightly infuscate. Head transverse, temples short; compound eyes not preserved; antenna five-segmented, with pedicel transverse, distinctly wider than flagellar base, flagellomere 1 thin, slightly widened apically, twice as long as wide, trapezoid, flagellomere 2 wider, together about as long as thick, unsegmented club. Pronotum not distinct, mesonotum only partially visible, with median longitudinal mesonotal sulcus and notauli well developed, meeting well before presumed position of scutellum (poorly preserved); mesopleuron with percurrent mesopleural groove; otherwise, thoracic structure obscure. Hind femora short. Forewing almost entirely invisible as preserved. Abdomen short, wide; ovipositor short, not exceeding abdominal apex, sheaths not distinct. Material. Holotype. Locality and age. Green River Formation, about eight miles south of De Beque, Colorado (Cockerell 1925), Ypresian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 25-26, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Cockerell, T. D. A. (1925) Fossil insects in the United States National Museum. Proceedings of the United States National Museum, 64, 1 - 15. [https: // www. biodiversitylibrary. org / item / 32410 # page / 7 / mode / 1 up]","Malaise, R. (1945) Tenthredinoidoa of south-eastern Asia with a general zoogeographical review. Opuscula Entomologica Suppl, 4 (288), 1 - 288. https: // doi. org / 10.1080 / 11035894509446460"]}

Leptostigma alaemacula n. sp. http://zoobank.org/ urn:lsid:zoobank.org:act: CB061CC9-BB17-4FED-B536-2A35796FCBBA Fig. 21. Holotype. SR 92-18 - 08 (Fig. 21). A forewing, collected by Carma Henry at Republic on September 25, 1992, in the collections of the SR. Diagnosis. Distinguished from other species of Leptostigma by a combination of forewing dark in cell 1r+2rm. Tentatively diagnostic: R+M shorter than 1-M; 2r-rs joins cell 3rm distinctly near its middle; 2r-m joins M scarcely basal to 2m-cu; anal cells widely separated by fusion of 1A and 2A+3A, without crossvein. Description. Holotype. Forewing length 14.2 mm, maximum width 4.1 mm, length/width 3.5; membrane lightly infuscate throughout except darker posterior to 1r-rs; pterostigma dark; R+M shorter than 1-M; maximum length of cell 2r shorter than 3r; 2r-rs shorter than Rs between 2r-m, 2r-rs, joining cell 3rm near its middle; 2r-m slightly curved, joins Rs notably basal to 2m-cu joining M, aligned with 2mcu; 1-M almost straight, sub-parallel to 1m-cu; 3r-m distinctly bowed toward base; cell 2mcu with M longer than 1m-cu; 2m-cu curved distinctly toward base; cu-a postfurcal; anal cells separated for distance greater than width of cell 2a, without crossvein between them. Remarks. Differs from L. brevilatum n. sp., L. longiclava n. sp., L. longipallidum n. sp., L. longitenebricum n. sp. by its forewing dark posterior to 1r-rs and 2r-m joins M close to 2m-cu. Differs from L. proxivena n. sp. and L. fasciatum n. sp. in which the anal cells are not separated by a fusion of 1A with 2A+3A. Material. Holotype. Etymology. From the Latin ala, meaning wing, and macula, spot, referring to the forewing spot. The epithet is a noun in apposition. Locality and age. Republic, Washington, United States of America, Tom Thumb Tuff Member of the Klondike Mountain Formation exposure A0307; mid-Ypresian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 22-23, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826

Archibald, S. Bruce, Rasnitsyn, Alexandr P. (2023): Cimbicidae (Hymenoptera, 'Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands. Zootaxa 5278 (1): 1-38, DOI: 10.11646/zootaxa.5278.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5278.1.1

Leptostigma fasciatum n. sp. http://zoobank.org/ urn:lsid:zoobank.org:act: 19480AA6-435A-490A-BB68-644C50F9BD32 Fig. 20. Holotype. GSC 142843 (Fig. 20). A mostly complete forewing, missing the apex and the basal portion of cell 1a. Collected by SBA at McAbee (GSC locality V-016800), May 16, 2001, in the collections of the GSC (Vancouver). Diagnosis. Distinguished from other species of Leptostigma by a combination of forewing mid-length with dark transverse fascia, including small part of cell 1mcu. Tentatively diagnostic: R+M shorter than 1-M, 2r-rs joins cell 3rm distinctly distal to its middle, 2r-m joins M scarcely basal to 2m-cu, anal cells meet at crossvein (1A and 2A+3A not fused). Description. Holotype. Forewing length: 11.1 mm, maximum width estimated 4.0 mm, length/width 2.8; membrane lightly infuscate throughout, darker posterior to base of pterostigma; pterostigma narrow, tapering, similar to width of C-R before pterostigma; R+M shorter than 1-M; maximum length of cell 2r shorter than 3r; 2r-rs shorter than Rs between 2r-m, 2r-rs; 2r-m curved, slightly sigmoidal, joins Rs notably basal to 2m-cu joins M, joins M scarcely behind 2m-cu; 1-M straight; cell 1mcu almost square; 3r-m slightly curved; cell 2mcu with M shorter than 1m-cu; 2m-cu curved toward base; cu-a interstitial; anal cells meet at crossvein, with adventitious oblique crossvein in cell 2a. Remarks. Differs from L. brevilatum n. sp., L. longiclava n. sp., L. longipallidum n. sp., L. longitenebricum n. sp. in having forewing with dark transverse fascia and 2r-m joining M close to 2m-cu. Differs from L. proxivena n. sp. by a more distal position of 2r-rs (distal to middle of cell 3rm), from L. alaemacula n. sp. by anal cells meeting at crossvein (1A and 2A+3A not fused). Material. Holotype. Etymology. From the Latin adjective fasciatus, meaning striped, referring to the forewing colouration of the species. Locality and age. McAbee, British Columbia, Canada, Hoodoo Face beds; mid-Ypresian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 21-22, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826

Allenbycimbex morrisae n. sp. http://zoobank.org/ urn:lsid:zoobank.org:act: 158DB20D-4B4D-41C3-B064-CCB1AB5237A9 Fig. 4. Material. Holotype RBCM.EH2007.002.0001a, b (part, counterpart). Mostly complete but somewhat deformed by geological shear forces, lacking head, fore and mid legs, hind tarsi, apical half of left wings, and apical half of abdomen; hind wings fragmentary and poorly preserved; wings lack visible colouration either originally or by preservation; collected by Muriel Morris, Allenby Formation; housed in the collections of the RBCM. Diagnosis. As for genus. Description. Length as preserved (stretched, see below; incomplete head and abdomen) 8.9 mm, apparently dark (preserved parts: hind tibiae, metasoma possibly paler). Temples somewhat inflated. Antennal club if correctly identified apparently short with truncated apex. Lateral pronotal lobes very wide, notauli apparently widely separated before scutellum (interpretation tentative). Hind coxae large, contiguous; femora not inflated, almost parallel-sided, shorter than abdomen wide; tibia thinner and much longer than femur. Forewing: length ca. 13.5 mm as preserved (distorted), the same in life if it was compressed in width, ca. 10 mm in life if it was stretched in length; maximum width: ca. 3.5 mm as preserved, about the same in life if it was stretched in length or ca. 4.25 if compressed in width; length/width: probably ca. 3.0; no membrane colouration detected (could be an artefact of preservation); pterostigma broad, distinctly wider than C-R basal to it; R+M longer than 1-M; 2r-rs not preserved; 2r-rm curved, joins Rs about same level as 2m-cu joins M, joins M behind 2m-cu for more than half length of 2r-m; 1-M almost straight; 1-M and 1m-cu diverging toward anterior; 3r-m distinctly bowed toward base; cell 2mcu with M shorter than 1m-cu; 2m-cu curved distinctly toward base; cu-a postfurcal; anal cell contracted for distance subequal to width (height) of 1a cell, with no crossvein; basal portion of 2A+3A sharply bent toward 1A, meets 1A at steep angle. Hind wing with m-cu not preserved, r-m and cu-a far distant (for much longer than in Cenocimbex and more similar to that characteristic of Leptostigma n. gen.); other details obscure. Abdomen with 3 segments preserved (apparently sterna 2–4) each narrower than thorax and some 3.5× as wide as long. Remarks. The fossil was distorted by geological shear forces during diagenesis, either stretching it along the plane of the length of its right forewing or compressing it 90° to that. These possibilities cannot be distinguished. As the forewings are about 90° to each other, we estimated its life shape by both graphically compressing the image along the length of the right forewing and by stretching it’s width so that elements of its venation are of equal length to those of its left wing. Etymology. The specific epithet is a patronymic formed from the surname of Muriel Morris, recognizing her contribution as collector and donor of the only known specimen., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on pages 7-8, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826

Subfamily Cenocimbicinae n. subfam. http://zoobank.org/ urn:lsid:zoobank.org:act: FA885793-D7D8-49A8-A00D-6C813ACBFC0F Type genus. Cenocimbex Nel, 2004. Diagnosis. Distinguished from all other Cimbicidae by forewing with 2r-m distinctly curved, meeting M distal of 2m-cu; cell 2r+pterostigma distinctly higher than cell 3r; and a combination of the following: eyes converging ventrally; antenna with 7–9 antennomeres, club 3-segmented; mesonotum with median longitudinal mesonotal sulcus and notauli distinct; metanotum short, with cenchri oval; tibial spur acute, long; first abdominal tergite with hind margin straight or weakly emarginated. Remarks. Cenocimbicinae differ from individual subfamilies of Cimbicidae by: from all by 2r-m as in diagnosis; from Abiinae, Cimbicinae by eyes converging ventrally [Cimbicinae: more or less parallel; Abiinae: converging dorsally], by at least one tibial spur acute, long (equal length of first tarsomere, one known, see F-774, tentative Leptostigma brevilatum) [Abiinae, Cimbicinae: blunt]; from Corynidinae, Pachylostictinae by 2r-m posterior insertion distal to cell 2mcu [Corynidinae, Pachylostictinae: proximal to], by cell 2r+pterostigma distinctly higher than cell 3r [not so], antenna with more than five antennomeres [five], club 3-segmented [non-segmented], metanotum short [long]; further from Corynidinae by mesonotum with median longitudinal mesonotal sulcus and notauli distinct [reduced or absent]; from some Abiinae and Cimbicinae by cenchri oval, from some Cimbicinae also by first abdominal tergite with hind margin straight or weakly emarginated [deeply emarginated]. Attribution of the Cenocimbicinae to the Tenthredinoidea is confident by 1-M meeting R and 2r-rs meeting Rs distally to 2r-m, and to the Cimbicidae by their clubbed antenna, and a unique forewing venation characteristic of the core cimbicid subfamilies Cimbicinae and Abiinae and found nowhere else in Hymenoptera: venation complete for Hymenoptera (except Sc, Rs between Rs+M and 1r-rs, and Rs bifurcation lost), characteristically narrow (low) cell 3r, and 2r-m crossvein meets M distal to 2m-cu. The long, curved to strongly curved crossvein 2r-m is distinct, occurring only rarely in extant Cimbicidae (very few Abiinae, cf. Vilhelmsen 2019, Fig. 16A; and Cimbicinae, cf. Vilhelmsen 2019, Fig. 16B) and in the Miocene of China (Abiinae: cf. Zhang et al. 1994, Fig. 121; Cimbicinae: cf. Zhang et al. 1994, Fig. 118), which we consider convergent. The new subfamily comprises the oldest known Cimbicidae, from the Selandian of Menat and less than ten million years later in the Ypresian Okanagan Highlands deposits of far-western North America (Archibald et al. 2011), a rather narrow interval of time but across a great distance of the Holarctic. Genera included. The type genus, Allenbycimbex n. gen. and Leptostigma n. gen. Locality and age. Menat, France, Selandian; Okanagan Highlands (McAbee,Allenby Formation, and Republic), far-western North America, Ypresian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on page 4, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Nel, A. (2004) New and poorly known Cenozoic sawflies of France (Hymenoptera, Tenthredinoidea, Pamphilioidea). Deutsche Entomologische Zeitschrift, 51, 253 - 269. https: // doi. org / 10.1002 / mmnd. 20040510208","Vilhelmsen, L. (2019) Giant sawflies and their kin: morphological phylogeny of Cimbicidae (Hymenoptera). Systematic Entomology, 44, 103 - 127. https: // doi. org / 10.1111 / syen. 12314","Zhang, J. - F., Sun, B. & Zhang, X. (1994) [Miocene insects and spiders from Shanwang, Shandong]. Shandong Science and Technology Publishing House, Jinan, 298 pp. [in Chinese with abstract in English]"]}

Leptostigma n. gen. http://zoobank.org/ urn:lsid:zoobank.org:act: 531F3657-F3D9-4A4D-86B7-1C146E74D627 Figs. 3A–D, 3G–K, 5–21. Type species. Leptostigma brevilatum n. sp. Diagnosis. Distinguished from Cenocimbex by lacking dark triangular sclerite on R basal to pterostigma reaching 1-Rs+M near 1m-cu (absent or undetectable being small, pale); wing narrower (length/width 2.8 or more); cell 3rm long (2r-m to 3r-m distinctly longer than length of 3r-m); hind wing with cu-a well before middle of cell mcu; distinguished from Allenbycimbex by pterostigma narrow, maximum height about equal to height to C to R basal to pterostigma. Species included. The type species and L. longiclava n. sp., L. longipallidum n. sp., L. longitenebricum n. sp., L. proxivena n. sp., L. fasciatum n. sp., and L. alaemacula n. sp. Etymology. From the Greek leptos, meaning thin, and stigma, for the pterostigma. The name is neuter (see Examples below Art. 30.1.2, ICZN 1999). Locality and age. Republic, Washington, United States of America, Tom Thumb Tuff Member of the Klondike Mountain Formation exposure A0307; and McAbee, British Columbia, Canada, Hoodoo Face beds; mid-Ypresian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on page 8, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["ICZN (1999) International Code of Zoological Nomenclature. 4 th Edition. The International Trust for Zoological Nomenclature, London, 306 pp. [https: // www. iczn. org / the-code / the-code-online /]"]}

Phenacoperga coloradensis (Cockerell, 1907) Fig. 26. Perga coloradensis Cockerell, 1907, p. 446. Phenacoperga coloradensis Cockerell, 1908, p. 113 –114, new combination. Holotype. UCM 4537 A rather complete female specimen missing most of the legs and the right wings. “ Perga ” coloradensis of Cockerell (1907) and see Phenacoperga of Cockerell (1908) and Rohwer (1908). Diagnosis. As for genus. Description. Holotype, female. Body: length ca. 27.7 mm; head, thorax, abdomen dark; antennae short, ca. 6.7 mm, 1.2 × head width (5.7 mm), dark, subclavate (very gradually widening apicad), with basal fragellomeres short (ratios of five apical segment lengths from base to apex, 1: 1.1: 0.8: 0.7: 1.7), apical one longer than two preapical combined; labrum large (ca. half as wide as separation of compound eyes); median longitudinal mesonotal sulcus, notauli present, well-developed. Forewing: membrane hyaline, length ca. 20.7 mm, maximum width estimated ca. 6.5 mm, length/width: ca. 3.3; pterostigma narrow, tapering, similar to width of C-R before pterostigma; R+M about same length as 1-M; maximum length of cell 2r shorter than 3r; 2r-rs longer than Rs between 2r-m, 2r-rs, joining Rs just basal to middle of cell 3rm; 2r-m nearly straight, short, joins M at 2m-cu; 1-M straight; cell 1mcu with M, 1m-cu divergent toward anterior margin; 3r-m straight; cell 2mcu with M longer than 1m-cu; 2m-cu apparently straight (faint); cu-a at branching of M, Cu; 2A complete, anal cells separated by crossvein 1a. Hind wing with m-cu much closer to 3r-m than to cu-a. Hind coxae small, apparently contiguous, hind femur and tibia thin. First abdominal tergum apparently not split medially, with hind margin rounded. Abdomen much wider than thorax, almost as long as forewing, ovipositor comparatively long (about one third as long as abdomen). Locality and age. Florissant Formation, Priabonian., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on page 29, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Cockerell, T. D. A. (1907) Some Old World types of insects in the Miocene of Colorado. Science, 26, 446 - 447. https: // doi. org / 10.1126 / science. 26.666.446","Cockerell, T. D. A. (1908) The fossil saw-fly Perga coloradensis. Science, 27, 113 - 114. https: // doi. org / 10.1126 / science. 27.681.113. b","Rohwer, S. A. (1908) On the Tenthredinoidea of the Florissant Shales. Bulletin of the American Museum of Natural History, 24, 521 - 530."]}

Genus Phenacoperga Cockerell, 1908 Fig. 26. Perga: Cockerell, 1907, p. 446, original description of P. coloradensis /misidentifiation of genus. Phenacoperga Cockerell, 1908, p. 113 –114. Type species. Perga coloradensis Cockerell, 1907. Emended diagnosis. Apparently distinct from other genera of Cimbicinae by antennae short (length ca. 1.2 × head width), subclavate, flagellomere 2 only twice as long as wide; 1-M, 1m-cu strongly divergent toward anterior margin; crossvein 2r-m slightly bent. Remarks. The forewing venation of Phenacoperga agrees with Cimbicinae as its M+Sc-R junction is well separated from that of Rs+M, and the posterior anal vein 2+3A is complete, anal cells joined by crossvein 1a. In the key to Asiatic Cimbicidae of Wei et al. (2012), which includes all genera of Cimbicinae but Labriocimbex Yan & Wei, 2019, Phenacoperga keys to couplet 12 with Pseudoclavellaria Schulz, 1906 and Leptocimbex Semenov, 1896 but differs from both in its huge size, short and stout antennae and 1-M and 1m-cu strongly divergent toward anterior margin. Phenacoperga additionally differs from Labriocimbex and Pseudoclavellaria by its apparent absence of a rich vestiture, from Pseudoclavellaria by its ill-defined antennal club, and from Leptocimbex by its robust stature., Published as part of Archibald, S. Bruce & Rasnitsyn, Alexandr P., 2023, Cimbicidae (Hymenoptera, ' Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands, pp. 1-38 in Zootaxa 5278 (1) on page 28, DOI: 10.11646/zootaxa.5278.1.1, http://zenodo.org/record/7894826, {"references":["Cockerell, T. D. A. (1908) The fossil saw-fly Perga coloradensis. Science, 27, 113 - 114. https: // doi. org / 10.1126 / science. 27.681.113. b","Cockerell, T. D. A. (1907) Some Old World types of insects in the Miocene of Colorado. Science, 26, 446 - 447. https: // doi. org / 10.1126 / science. 26.666.446","Yan, Y., Niu, G., Zhang, Y., Ren, Q., Du, S., Lan, B. & Wei, M. (2019) Complete mitochondrial genome sequence of Labriocimbex sinicus, a new genus and new species of Cimbicidae (Hymenoptera) from China. PeerJ, 7, e 7853 https: // doi. org / 10.7717 / peerj. 7853","Schulz, W. A. (1906) Strandgut. Spolia Hymenopterologica, [1906], 76 - 269. https: // doi. org / 10.5962 / bhl. title. 59757","Semenov, A. [Semenow, A.] (1896) De Tenthredinidarum genere novo Clavellariae Oliv. proximo. Ezhegodnik \" Zoologicheskago Muzeja Imperatorskoj Akademii Nauk \", 1, 95 - 104."]}