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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. 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(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"]}

Diagnostic key for the Kathistaceae 1. Sexual morph present with sporodiomata...................................................................................................................................... 10 - Asexual morph present with sporodochium..................................................................................................................................... 2 2. The production of microconidia and macroconidia appendages................................................................................ T. cavernosum - Microconidia produced at conidiogenous locus................................................................................................................................3. 3. Phialide tips echinulate and pigmented........................................................................................................................... T. coronata - Phialide tips flat and unpigmented.................................................................................................................................................. 4. 4. Phialides greater than 160 μm in length................................................................................................................... T. longiphialidis - Phialides less than 120 μm in length............................................................................................................................................... 5. 5. Conidia are large; dimensions ~ 8 μm × 4 μm............................................................................................................ T. macrospora - Conidia are small; dimensions ~4 μm × 2 μm................................................................................................................................ 6. 6. Large rhomboid or ellipsoid sporodochia; 5–6 mm in diameter.............................................................................. T. rhombicarpa - Circular or ellipsoid sporodochia; less than 1 mm in diameter....................................................................................................... 7. 7. Maximum of 5 conidia per collarette................................................................................................................................. T. snyderi - Minimum of 8 conidia per collarette............................................................................................................................................... 8. 8. Phialides tips exposed at surface of sporodochium and terminate in flaps; phialides are fertile and not arranged into compartments.......................................................................................................................................... T. hexasporodochia sp. nov. - Phialide tips covered at surface of sporodochium by superficial epihymenium; phialides arranged into sterile and fertile compartments.................................................................................................................................................................................. 9. 9. Sporodochia exhibit a beak-like pycnidium and ostiole and are ellipsoid in shape; conidiogenous locus ~15 μm from base.................................................................................................................................................................................................... M. crustosa - Sporodochia lack pycnidia and ostiole; sporodochia are stellate or ellipsoid in shape; conidiogenous locus ~ 50 μm from base................................................................................................................................................................................................ M. silvestri 10. 4 ascospores produced in ascus........................................................................................................................... Kathistes fimbriata - 8 ascospores produced in ascus........................................................................................................................................................11 11. Ascospores develop with 3 septa........................................................................................................................ Kathistes calyculata - Ascospores develop with 5 septa................................................................................................................ Kathistes analemmoides

Species: Termitaria hexasporodochia Wilson, Emam, Davis, Barden, Hall, Ware. sp. nov. Mycobank ID: MB832822 Newark Museum Herbarium, Newark, NJ (NEMU) American Museum of Natural History, NYC, NY (AMNH) Type Specimens Locality: South America, Guyana: Rupununi River Region, Karanambu Ranch, Capuchin trail, elevation: 100km, 03°44.85’ N,059°19.13’ W, tropical rainforest surrounded by open savannah, on host termite in an arboreal soil mound, 11, January 2016, Collected by M. Wilson, J. Ware, P. Barden, S. George. L. Johnson, S.T. Mafla-Mills. On Amitermes sp., preserved in ethanol. Deposited at AMNH and registered at NEMU Herbarium (NEMU), labeled: South America, Guyana: Rupununi River Region, Karanambu Ranch, Capuchin trail, elevation: 100km, 03°44.85’ N, 059°19.13’ W, collected 11, January 2016, (holotype: T1-P1S7S3!, isotypes: T2-P1S7S3!, T2-P2S7S3!). Diagnosis— Termitaria hexasporodochia sp. nov. exhibits a unique phenotype, exclusively forming six moderately sized black rimmed, elliptical sporodochia on the ventral abdominal segments (4–6) of its host. Termitaria hexasporodochia sp.nov. exhibits8–9conidia per collarette,is 116–120μm thick,elliptical in shape, has a conidiogenesis zone location 20μm–25μm from the sporodochium base, and thick-walled, lobate haustoria. The dimensions of the sporodochia range from 256 μm–609μm to 102–218 μm, the phialides are 110–115 μm in length, and the expiculum is the only pigmented region noted. Description of the holotype —Entomogenous: Sporodochium thallus is 116 μm thick, 256 μm long, 102 μm wide, and ellipsoid in shape. At the basal layer, haustoria are initially formed from thick-walled haustorial mother cells at the base of the sporodochium and appear thick and lobular. Haustoria range from 19–23μm in length, and 1.7–2.4 μm in width. Superficial to this layer is the sporogenous hymenium layer composed of a mass of tight columnar phialides 110–115 μm thick. Each individual phialide ranges from 1.5–2.2 μm in diameter, terminating in two flat rounded tips. Endogenously-formed conidia originate from long rod-shaped conidiogenous cells at the conidiogenous locus, located 20–25 μm from the base of each phialide. Conidial spores are rectangular and catenate, 1–1.3 μm in width and 3–4 μm in length. 8–9 spores were found in each collarette and 11–13 in each phialide. Spores break off simultaneously as they reach the sporodochium surface. The surface of each sporodochial lesion appears perforated, bearing a hexagonal pattern referred to as “textura angularis”, with pores being larger in diameter along the rim preceding the sterile expiculum. The expiculum forms a smooth black crust with no apparent openings. The holotype selected for description was one of the six lesions found infesting the 4 th ventral abdominal segment of an Amitermes termite worker T1-S7S3!. Etymology— Termitaria is the established generic name, and hexasporodochia is an adjective referring to the hexad arrangement of sporodochia exhibited on the type-host, Amitermes. Ecology and host species— Amitermes sp. found to be infested with Termitaria were collected from large arboreal mounds constructed on sandpaper trees (Curatella americana) growing at the interface between open savannah and rainforest (Fig. 2). The mounds were found during the dry season (January) in regions surrounding the Rupununi River that flood during the annual wet season (May-August). It was found only on worker caste termite hosts, never on soldiers or reproductives. Differential diagnoses— Termitaria hexasporodochia sp. nov. exhibits a unique arrangement phenotype, forming six moderately-sized, black-rimmed, elliptical sporodochia on the ventral abdominal segments (4–6) of its host. This fungus belongs to the genus Termitaria and is readily distinguished from two described members of the genus Mattirolella. It lacks a major generic character shared amongst Mattirolella, the sterile hyphae separating fertile hyphae in the hymenium is not present in the new species. The maximum lengths of T. hexasporodochia sp. nov. sporodochia are less than half the size of those described for Australian species, T. macrospora and T. rhombicarpa (Table 1; Kimbrough & Lenz 1982). Additionally, T. rhombicarpa can be distinguished by the rhomboid shaped sporodochium it typically forms on the host as opposed to the elliptical/circular lesion formed on T. hexasporodochia sp. nov., shorter phialide lengths (Table 1, 100–105 μm vs. 110–115μm), longer conidiogenous locus (Table 1. 30–35 μm vs. 20–15 μm from phialide base) and haustoria penetrating twice as deep into the host cuticle (TABLE 1. 50–55 μm vs. 20–23 μm). T. hexasporodochia sp. nov. spores are easily distinguished from those produced by T. macrospora. As its name suggests T. macrospora produces 4–5 massive spores per collarette (app 3.8–9.1 μm) whereas T. hexasporodochia sp. nov. produces 8–9 spores per collarette (app 1–1.3 × 3–4 μm). Another Australian species, T. longiphialidis, is best identified by its small circular sporodochia and having the longest phialides of any Termitaria species (Table 1. 160–180 μm vs. 110–115 μm; Kimbrough & Lenz 1982). T. hexasporodochia sp. nov. is readily distinguished from the commonly described T. snyderi, by phialide length. T. hexasporodochia sp. nov. possesses a much thicker sporodochium, with phialides double the length of those found in T. snyderi (Table 1, 110–115 μm vs. 50–60 μm). T. coronata can be distinguished from all Termitaria species, T. hexasporodochia sp. nov. included, in the appearance of its sporodochium, with its echinulate surface and the high position of its conidiogenous locus (Table 1, 50–60 μm vs. 20–25 μm). Ultrastructure of T. hexasporodochia sp. nov. —The complete sporodochium (Fig.4) is composed of many tightly aligned vertical columns. We group the sporodochia layers into three major regions (Fig.5A); the basal region (Fig. 5) most closely appressed to the insect cuticle, the sporogenous hymenial region (Fig. 6), and the upper region (Fig.7) filled with conidia and phialides terminating into flaps. The basal region —This region of the sporodochium is approximately 4–5 rows and 8.5–12.5 μm thick. It is comprised of haustorial mother cells that give rise to a subcuticular layer of haustoria that penetrates the host (Fig. 5). The thin-walled cells composing the upper layer of the basal region have been referred to in previous studies as the “subhymenial layer” and gives rise apically to the hymenial phialides (Kimbrough and Thorne 1982). Haustoria aggregate towards and penetrate the host via tetra ocular channels in its cuticle. We observe 12+ major penetration points between the fungal body and its host (Fig.5 B), extending below the cuticle to thick-walled lobular haustoria ranging from 19–23 μm in length and 1.7–2.5 μm in width. We do not refer to this “haustorial region” as an internal region within the sporodochium because it lays below the cuticular layer of its host (Fig.6E–F). Additionally, its penetration into the host cuticle does not change the classification of this fungus as an ectoparasite, in that it does not invade the host cytoplasm. Each sporodochia appears to be formed from major infestation sites with cells growing upward from the basal layer, with approximately 160–240 openings per sporodochium (size dependent) that appear dark in photomicrographs (Fig.5C). The sporogenous hymenium region —This region consists of a thick hymenium of tightly appressed phialides (Fig.6A), comprised of long rod-shaped conidiogenous cells 14–22 μm in length, which differentiate into asexual conidia at a fixed conidiogenous locus (Fig 6B–C). The phialides are approximately 1.5–1.2 μm in diameter and the conidiogenous locus (Fig.6D), the initial point of asexual spore differentiation, occurs 20–25μm, from the base of each hymenial phialide. Each collarette contains 8–9 rectangular, catenate conidia (Fig.7B–C), and approximately 11–13 spores can be found in the phialide during this time. Under high-definition microscopy, the internal surface of the hymenial phialides are coated with a dense mat of minute filaments that appear to be restricted to the upper 3/4th of the secondary canals (Fig.7A). The upper region — At the superficial level of the sporodochia, a dark, peripheral expiculum surrounds a region resembling a hexagonal honeycomb (Fig.7A,C). This pattern is formed by fields of terminating phialides (Fig.3B) of elongate tubes referred to as textura angularis, the apices of which are bivalved (Fig 7B–D). The apical valves are formed by two isosceles trapezoidal flaps that fit closely together to form a large circular pad of thousands of hexagons (Fig.7D). Visible from confocal stack images just below the pad surface, each sporodochia appears densely populated with hexagonal pores, with the conidial spores visible within each tubular hymenial channel leading to the apical pore valves (Fig.7B). On average, a sporodochia contains approximately 12,000 –14,000 phialides. Overall, the most striking feature of Termitaria hexasporodochia sp. nov. is apparent in the arrangement and ultrastructure of the six moderately-sized, black-rimmed, elliptical sporodochia it forms on its termite host. Of all members of the family Kathistaceae, the formation of sporodochia on termites is diagnostic for members of the genera Termitaria and Mattirolella, hence why we focus on those two genera in assessing the new species. Termitariopsis is the only remaining member of Kathistaceae which forms sporodochia, although it does not use termites as a host and exhibits sporodochial features absent in the other two genera (Table 2a). Kathistes genera do not form sporodochia and are described instead off of their sexual form, whereas Termitaria hexasporodochia sp. nov. is described by its asexual sporodochia form. The genus Ectomyces calotermi, described on termites to form sporodochial lesions, has been synonymized with Termitaria snyderi (Tate 1928), a species described in this article. No further examination is needed on this genus as its synonym T. snyderi is examined here. We place the new species in Termitaria because it lacks the defining features of Mattirolella: sterile hyphae interspersed with fertile hyphae and an epihymenium (Table 2a). The new species can be differentiated from other Termitaria species by sporodochial length (T. macrospora and T. rhombicarpa), sporodochial shape (T. rhombicarpa), phialide lengths (T. longiphialidis, T. snyderi), location of conidiogenous locus, spore size, and haustorial depth (Table 2a)., Published as part of Wilson, Megan M., Emam, Amany, Davis, Steven R., Hall, Gene, Barden, Phillip & Ware, Jessica L., 2023, Description of a novel termite ectoparasite, Termitaria hexasporodochia sp. nov. (Kathistaceae), presenting an unusual six-sectioned infestation, and a key to the fungal family Kathistaceae, pp. 106-124 in Phytotaxa 591 (2) on pages 111-113, DOI: 10.11646/phytotaxa.591.2.3, http://zenodo.org/record/7797494, {"references":["Kimbrough, J. W. & Lenz, M. (1982) New species of Termitaria (Termitariales: Deuteromycetes) on Australian termites (Isoptera). Botanical Gazette 143: 262 - 272. [https: // www. jstor. org / stable / 2474717]","Khan, S. R. & Kimbrough, J. W. (1974 a) Taxonomic position of Termitaria and Mattirolella (entomogenous Deuteromycetes). American Journal of Botany 61: 395 - 399. https: // doi. org / 10.2307 / 2441806","Blackwell, M. (1980) New records of termite-infesting fungi. Invertebrate Pathology 35: 101 - 104. https: // doi. org / 10.1016 / 0022 - 2011 (80) 90093 - 2","Blackwell, M. & Rossi, W. (1986) Biogeography of fungal ectoparasites of termites. Mycotaxon 25: 581 - 601.","Colla, S. (1929) Su alcuni funghi parassiti delle Termiti. Bolletino del Laboratorio di Zoologia Portici 22: 39 - 48.","Rossi, W. R. & Rossi, M. G. C. (1977 b) Due Nuove Specie di Dimeromyces (Laboulbeniales). Rivista di Parassitologia 38: 111 - 113.","Tate, P. (1927) On Ectomyces calotermi n. g., n. s p., an ascomycete parasitic on Calotermes samoanus Holmgren (Isoptera, Porotermitidae). Parasitology 19: 54 - 60.","Kimbrough, J. W. & Thorne, B. L. (1982) Structure and development of Mattirolella crutstosa (Termitariales, Deuteromycetes) on Panamanian termites. Mycologia 73: 201 - 209. https: // doi. org / 10.1080 / 00275514.1982.12021492","Malloch, D. & Blackwell, M. (1990) Kathistes, a new genus of pleomorphic ascomycetes. Canadian Journal of Botany 68: 1712 - 1721.","Tate, P. (1928) Notes on the Genera Ectomyces and Termitaria, Fungi Parasitic on Termites. Parasitology 20 (1): 77 - 78. https: // doi. org / 10.1017 / S 003118200001146 X"]}

In recent decades, a lack of available knowledge about the magnitude, identity and distribution of biodiversity has given way to a taxonomic impediment where species are not being described as fast as the rate of extinction. Using Machine Learning methods based on seven different algorithms (LR, CART, KNN, GNB, LDA, SVM and RFC) we have created an automatic identification approach for odonate genera, through images of wing contours. The training population is composed of the collected specimens that have been digitized in the framework of the NSF funded Odomatic and TOWD projects. Each contour was pre-processed, and 80 coefficients were extracted for each specimen. These form a database with 4656 rows and 80 columns, which was divided into 70% for training and 30% for testing the classifiers. The classifier with the best performance was a Linear Discriminant Analysis (LDA), which discriminated the highest number of classes (100) with an accuracy value of 0.7337, precision of 0.75, recall of 0.73 and a F1 score of 0.73. Additionally, two main confusion groups are reported, among genera within the suborders of Anisoptera and Zygoptera. These confusion groups suggest a need to include other morphological characters that complement the wing information used for the classification of these groups thereby improving accuracy of classification. Likewise, the findings of this work open the door to the application of machine learning methods for the identification of species in Odonata and in insects more broadly which would potentially reduce the impact of the taxonomic impediment.

Danowhetaksa rusti Simonsen, Ware, & Archibald, new species Figure 2 Material. Holotype [MM-13418]: an isolated wing preserved in a concretion block, deposited in Museum Mors (Mo-clay Museum), collected by Henrik Madsen, October 31, 1993, Fur Stolleklint. The specimen was collected from a hardened bed within the upper 1.5 m of the Stolleklint clay near ash-layers -33 and -34. Description. Holotype wing. Arculus to distal end of pterostigma: 26.7 mm. Nodus to distal end of pterostigma: 18.9 mm. Arculus to basal end of pterostigma: 22.6 mm. Nodus to basal end of pterostigma: 14.8 mm. Width: 8.8 mm. Pterostigma dark, 5.2 mm in length, ca. 8 times longer than wide, subtends numerous cells, eight crossveins detected by preservation, surely more. Wing hyaline with transverse dark fascia, as in diagnosis, ca. 1.5 times length of pterostigma. Postnodal and postsubnodal spaces poorly preserved, but only one pair of crossveins appears aligned. IR1 very poorly preserved, origin probably zigzagged. RP2 originates 8 cells distal to subnodus. IR2 originates ca. 8/10 distance arculus to nodus, preserved part of IR2 close to linear. RP3-4 originates ca. 4/10 distance arculus to nodus, preserved part of RP3-4 close to linear. Preserved part of MA linear through fascia. Preserved part of MP linear (terminus missing). CuA linear from origin to terminus on wing margin, basal 2/3 subparallel to MA, then curving sharper to wing margin, terminates just to mid-wing beyond fascia. MA-CuA space two cells wide where CuA starts curving away, widening to at least four cells well before estimated terminus at wing margin. CuA-A space moderately well preserved, very broad, at least 5 cells wide. Quadrangle sub-trapezoid broadest distally, approximately 1.3x longer than wide. Ax0 present, synsclerotised with wing base. Arculus close to, slightly distal of Ax1. Ax2 partly preserved, ca. fifth distance arculus to nodus, just distal to quadrangle. Diagnosis. Distinguished from D. birgitteae as in its diagnosis, above. Deposit and age. Stolleklint clay, ��lst Formation, Stolleklint, Fur, Denmark; earliest Ypresian. Etymology. An eponym formed by the surname of the German paleoentomologist Jes Rust, whose extensive work has greatly increased our knowledge of mo-clay insects. Remarks. Garrouste & Nel (2015) described the monobasic Pseudostenolestidae from the latest Ypresian at Messel, Germany (Pseudostenolestes bechlyi Garrouste & Nel) and discussed ways in which it resembles the Dysagrionidae and Sieblosiidae (Cephalozygoptera). It shares a distinctive quadrangle shape with the Whetwhetaksidae, which the Dysagrionidae and Sieblosiidae also possess. Notably, the arculus of P. bechlyi is positioned near Ax1 as in the Whetwhetaksidae. In P. bechlyi, the arculus and Ax1 are opposite, while in Whetwhetaksidae Ax 1 is just basal to the arculus���slightly closer to it in Danowhetaksa than in Whetwhetaksa. The pterostigma is also long in P. bechlyi, although not as long as in the Whetwhetaksidae. Unlike the Whetwhetaksidae, however, it possesses the oblique vein ���O���, shared with the Sieblosiidae, and unlike the Whetwhetaksidae, Sieblosiidae, and Dysagrionidae, it has a very short petiole. Pseudostenolestes differs from the Zygoptera, Cephalozygoptera, and Whetwhetaksidae by its distinctive, strong vein Cuab originating at the middle posterior of the subquadrangle and directed towards the wing base, by which they assign it to the Isophlebioptera, previously only known from the Mesozoic. In most Zygoptera, Ax0 is absent or obscured by sclerotization at the wing base (Bechly 1996, and see Rehn, 2003). It is present without associated sclerotization in Whetwhetaksa and in Cephalozygoptera (Archibald et al. 2021). In D. rusti n. sp. it is present and associated with sclerotization basally, but not obscured by it., Published as part of Simonsen, Thomas J., Archibald, S. Bruce, Rasmussen, Jan A., Sylvestersen, Ren�� L., Olsen, Kent & Ware, Jessica L., 2022, Danowhetaksa gen. nov. with two species from the early Eocene ��lst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera), pp. 586-592 in Zootaxa 5099 (5) on pages 589-590, DOI: 10.11646/zootaxa.5099.5.5, http://zenodo.org/record/6110289, {"references":["Garrouste, R. & Nel, A. (2015) New Eocene damselflies and the first Cenozoic damseldragonfly of the isophlebiopteran lineage (Insecta: Odonata). Zootaxa, 4028 (3), 354 - 366. https: // doi. org / 10.11646 / zootaxa. 4028.3.2","Bechly, G. (1996) Morphologische Untersuchungen am Flu ¨ gelge der der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata) unter besonderer Beru ¨ chsichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, B blingen, Special Volume, 2, 1 - 402.","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","Archibald, S. B., Cannings, R. A., Erickson, R. J., Bybee, S. M. & Mathewes, R. W. (2021) The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America. Zootaxa, 4934 (1), 1 - 133. https: // doi. org / 10.11646 / zootaxa. 4934.1.1"]}

Danowhetaksa Simonsen, Ware & Archibald, new genus Diagnosis. Most easily distinguished from Whetwhetaksa by: 1, pterostigma length ca. 7 times width [ca. 10 times]; 2, dark fascia mid-wing, basal wing hyaline [colouration extends to wing base where known]; 3, origin of IR2 closer to nodus than to origin of R3-4 [closer to origin of R3-4], at least three crossveins between them [one]; 4, MP, CuA more widely separated distally [maximum ca. 10 cells at margin], not subparallel [maximum two cells, subparallel]. Type and included species. Type species, Danowhetaksa birgitteae n. sp. here designated; other included species, Danowhetaksa rusti n. sp. Etymology. The genus name is formed from the prefix ���Dano-��� referring to Denmark, and the suffix ���-whetaksa��� referring to the Whetwhetaksidae. Gender: feminine., Published as part of Simonsen, Thomas J., Archibald, S. Bruce, Rasmussen, Jan A., Sylvestersen, Ren�� L., Olsen, Kent & Ware, Jessica L., 2022, Danowhetaksa gen. nov. with two species from the early Eocene ��lst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera), pp. 586-592 in Zootaxa 5099 (5) on pages 587-588, DOI: 10.11646/zootaxa.5099.5.5, http://zenodo.org/record/6110289

Danowhetaksa birgitteae Simonsen, Ware & Archibald, new species Figure 1 Material. Holotype [FUM-M-17515]: an isolated wing preserved in a concretion block; the part is missing the basalmost and apical-posterior portions, and the counterpart is missing the apical-most portion including the pterostigma; collected by Birgitte Munk, 1996, Fur Stolleklint; deposited in the Fur Museum. Description. Holotype wing. Length, arculus to distal end of pterostigma: 28.0 mm; nodus to distal end of pterostigma: 20.3 mm; arculus to base of pterostigma: max 23.4 mm; nodus to base of pterostigma: max 16.2 mm; width: 9.3 mm. Pterostigma dark (damaged basally), preserved part approximately 5 mm in length, highly elongate, at least seven times longer than wide, subtends numerous cells, five crossveins detected by preservation, surely many more. Membrane with a transverse, sub-central dark fascia, slightly narrower than length of pterostigma, otherwise hyaline. Twenty-three crossveins preserved in postnodal space, 22 in postsubnodal space, only pair aligned. IR1 very poorly preserved, origin probably slightly zigzagged, seven cells distal to origin of RP2. RP2 originates seven cells distal to subnodus. IR2 originates approximately 8.5/10 distance arculus to nodus, preserved part of IR2 close to linear. RP3-4 originates approximately 4/10 distance arculus to nodus, preserved part of RP3-4 close to linear. MA linear in basal 1/3, starts zigzagging slightly about basal margin of darkened wing band, poorly preserved beyond band. MP linear from origin to terminus at wing margin, slightly curved. CuA linear from origin to terminus on wing margin, basal 2/3 subparallel to MA, then curving sharper to wing margin, terminates basal to mid-wing. MA-CuA space two cells wide where CuA starts curving away, widening to at least six cells at wing margin. CuA-A space well preserved, very broad, at least five cells wide. Quadrangle sub-trapezoid, broadest distally, approximately 1.3 times longer than wide. Anterior wing margin not preserved basal to arculus, so Ax1 not preserved. Ax2 approximately 2/10 distance arculus to nodus, just distal to anterodistal corner of quadrangle. Diagnosis. Distinguished from D. rusti by any of: 1, RP1-2���IR2 space one cell wide, RP2���IR2 space to origin of IR1 (not preserved beyond this) [RP1-2���IR2 space becomes two cells wide ca. two cells basal origin of RP2]; 2, IR2���RP3-4 space becomes two cells wide ca. mid-way between origins of RP2, IR1 [ca. origin of IR2]; 3, seven crossveins in R1+2���IR2 space between nodus and origin of RP2 [ca. 9]. Further separated by colouration (sex unknown): narrow, dark fascia mid-wing [almost twice as wide], basally starting ca. five cells distal to nodus, includes origin of RP2, ends slightly distal to termination of CuA [anterior to R1+2 starts at nodus, posterior R1+2 just basal origin of IR2, ends on posterior margin just basal to termination of CuA, on anterior margin slightly more distal]. Deposit and age. Stolleklint clay, ��lst Formation, Stolleklint, Fur, Denmark; earliest Ypresian. Etymology. An eponym formed from the given name of Birgitte Munk, who found and donated the holotype, recognising her contribution. Remarks. Colouration is provided as supplementary in the diagnosis, as it is unknown if there are differences in colouration due to sexual dimorphism, variation between forewings and hind wings, or polymorphism in Whetwhetaksidae. Although the wing base is not preserved, we estimate CuA to terminate on the margin just over half wing length., Published as part of Simonsen, Thomas J., Archibald, S. Bruce, Rasmussen, Jan A., Sylvestersen, Ren�� L., Olsen, Kent & Ware, Jessica L., 2022, Danowhetaksa gen. nov. with two species from the early Eocene ��lst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera), pp. 586-592 in Zootaxa 5099 (5) on pages 588-589, DOI: 10.11646/zootaxa.5099.5.5, http://zenodo.org/record/6110289

Family Whetwhetaksidae Archibald & Cannings Remarks. We emend the diagnosis of Whetwhetaksidae of Archibald et al. (2021) based on the fossils described here by the following: character state 1, pterostigma at least seven times as long as it is wide (this was ten times width in Archibald et al. 2021), and remove character state 10, base to nodus percent wing length, as a synapomorphy for this group as this region is shorter in Danowhetaksa and does not now distinguish Whetwhetaksidae from Dysagrionidae or Sieblosiidae., Published as part of Simonsen, Thomas J., Archibald, S. Bruce, Rasmussen, Jan A., Sylvestersen, Ren�� L., Olsen, Kent & Ware, Jessica L., 2022, Danowhetaksa gen. nov. with two species from the early Eocene ��lst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera), pp. 586-592 in Zootaxa 5099 (5) on page 587, DOI: 10.11646/zootaxa.5099.5.5, http://zenodo.org/record/6110289, {"references":["Archibald, S. B., Cannings, R. A., Erickson, R. J., Bybee, S. M. & Mathewes, R. W. (2021) The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America. Zootaxa, 4934 (1), 1 - 133. https: // doi. org / 10.11646 / zootaxa. 4934.1.1"]}