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Ypopteron new genus (Fig 4.) Diagnosis. Tegmen of Ypopteron most easily separated from those of other Palaeorehniidae as in Table 1. Type and only species. Y. nicola. Description. As for its only species, see below. Etymology. The genus name refers to the Ypresian age, and ��� opteron, from the Greek -pteron (���������), meaning wing. Gender, neuter. Distribution and age. As for its only species, below., Published as part of Archibald, S. Bruce, Gu, Jun-Jie & Mathewes, Rolf W., 2022, The Palaeorehniidae (Orthoptera, Ensifera, " Zeuneropterinae "), and new taxa from the early Eocene Okanagan Highlands, western North America, pp. 559-572 in Zootaxa 5100 (4) on page 566, DOI: 10.11646/zootaxa.5100.4.6, http://zenodo.org/record/6225002

Republicopteron new genus (Figs. 5–7) Type and only species. Republicopteron douseae. Diagnosis. Tegmen of Republicopteron most easily separated from those of other Palaeorehniidae as in Table 1. Description. As in species description. Etymology. The genus name is a toponym referring to the town of Republic, Washington, and – opteron, from the Greek -pteron (πτερόν), meaning wing. Gender, neuter. Distribution and age. As for its only species, below.

Republicopteron douseae new species (Figs. 5���7) Diagnosis. As for genus. Description. Holotype tegmen. As in diagnosis and the following. Darkly infuscate throughout wherever preserved (darkness differing among specimens presumably by preservation) with some light spots between veins, crossveins as in P. maculata, but not as many or as distinct (perhaps preservation?). ScP with eight accessory branches preserved, surely more in missing basal region; no stridulatory file detected on CuPb. Type specimens. Holotype. SR 00-04-06 part, no counterpart (Fig. 5), a well-preserved anterior basal-mid region of a tegmen, precostal region not preserved, with a fragmentary portion of a second wing closely associated and a beetle preserved between fragments; deposited in SR; collected by Cassie Douse at Republic locality B4131, May 1, 2000, labelled: Holotype, Republicopteron douseae Archibald, Gu, and Mathewes. Paratype 1: SRUI 09- 95 - 73, (Fig. 6), collected by Michael Santino at Republic exposure A0307, April 19, 2009; labelled Republicopteron douseae Archibald, Gu, and Mathewes Paratype 1. Paratype 2: SR 21-005-001, (Fig. 7), collected by Gregg Wilson at Republic exposure B4131, May 14, 2014; labelled Republicopteron douseae Archibald, Gu, and Mathewes Paratype 2. Locality and age. Republic, Washington, USA; Tom Thumb Tuff Member of the Klondike Mountain Formation exposures B4131, 51.18 0.09 Ma. old and A0307, 51.45 0.12 Ma. old, mid Ypresian. Etymology. The specific epithet is a patronym formed from the surname of Cassie Douse, collector and donor of the holotype., Published as part of Archibald, S. Bruce, Gu, Jun-Jie & Mathewes, Rolf W., 2022, The Palaeorehniidae (Orthoptera, Ensifera, " Zeuneropterinae "), and new taxa from the early Eocene Okanagan Highlands, western North America, pp. 559-572 in Zootaxa 5100 (4) on pages 568-569, DOI: 10.11646/zootaxa.5100.4.6, http://zenodo.org/record/6225002

Family Palaeorehniidae Zeuner stat. nov. Palaeorehnia in the Hagloidea, Zeuneropterinae in the Stenopelmatoidea. Cockerell (1908) discussed the similarities of Palaeorehnia Cockerell to other taxa but did not assign it to a higher taxon within the Ensifera, nor did he publish a drawing of it, only low-resolution photographs (1909). Zeuner (1937) grouped Palaeorehnia and Jurassobatea Zeuner (Jurassic of Germany) as the Palaeorehniinae, a subfamily of the Gryllacrididae. Palaeorehnia included P. maculata (Scudder) from the Priabonian shale at Florissant, Colorado, USA (Scudder 1890; Cockerell 1908, 1909; Kevan and Wighton 1983) and P. scotica. Sharov (1962) treated P. maculata and Jurassobatea as Haglidae incertae sedis and erected the genus Zeuneroptera Sharov for P. scotica, maintaining the subfamily name Palaeorehniinae. As it no longer contained Palaeorehnia, Kevan and Wighton (1983) proposed the replacement name Zeuneropterinae (in Gryllacrididae), consisting only of Zeuneroptera. They assigned their new genus Albertoilus Kevan and Wighton to the Prophalangopsidae (Hagloidea) and suggested that P. maculata belongs to it as well. Zeuner, Sharov, Ragge (1955) and others in much of the Twentieth Century had differing arrangements of the superfamilies of Ensifera and their compositions, understanding the relationships of Zeuneroptera and Palaeorehnia to each other and within the Ensifera in a variety of ways. For brief reviews of the history of this thought, e. g., see Kevan and Wighton (1981, 1983) and Gorochov (2001). The current generally accepted view has Palaeorehnia in the Hagloidea and Zeuneropterinae (as Zeuneroptera and Albertoilus) in the Stenopelmatoidea (Gorochov 1995, but see B��thoux 2012, who makes an argument for Zeuneroptera in the Prophalangopsidae). Gorochov (2001) thought that the Zeuneropterinae might be close to the Anostostomatidae (Mimnermidae), especially its subfamily Cratomelinae, although with uncertainty as to it belonging the Stenopelmatoidea. In Gorochov���s diagnosis of the Stenopelmatoidea (1995, page 186), CuA+CuPa��, CuPa��, CuPb, and 1A are long and at a low angle to the posterior margin (���parallelization���), ending in the distal quarter of the wing as is reconstructed in the Zeuneropterinae. This is contra his Hagloidea concept (1995, page 107) where these four veins meet the posterior margin farther from the wing apex at a steeper angle (and see Gorochov 1988, 2001) as in the reconstruction of Cockerell���s P. maculata wing of Zeuner (1939: plate 2; plate 22, Fig. 1; plate 25, Fig. 1). Reassessment of superfamily assignments. The angles and ending points of CuA+CuPa��, CuPa��, CuPb, and 1A in Palaeorehnia, Zeuneroptera and Albertoilus, have, however, been estimated based on reconstructions of partial wings, none of which completely preserves those veins to their ends. To evaluate the assumption that these veins conform with Gorochov���s Hagloidea in Palaeorehnia, we examined modern high-resolution photographs of the part and counterpart of Cockerell���s holotype (Fig. 1A���1C). In these, the line thought to be a portion of the posterior wing margin can now be seen to be a part of a displaced, upturned vein in a folded section as in the similarly folded basal posterior region of the Ypopteron nicola type specimen (Fig. 4). Zeuner���s reconstruction (1939) then incorrectly rotates the wing clockwise, increasing the angles of these four veins to the non-existent portion of the posterior margin, therefore, ending too basally on the wing margin for Palaeorehnia to be associated with the Stenopelmatoidea (Fig. 1D). While there is not a sufficient portion of the posterior margin preserved to act as a landmark with which to orient the wing, it should be rotated counterclockwise by some unknown amount from Zeuner���s reconstruction and from that in Fig. 1E, and it is equally likely that the angles of these veins match those in Zeuneroptera and Albertoilus. This interpretation is also suggested by a new, high-resolution photograph of a second, unpublished fossil tegmen in the University of Colorado (Boulder) collections (UCM-18757: Fig 1E, F) labelled ��� Palaeorehnia maculata Ckll (apparently) Florissant T. Duce���. It is quite damaged and is somewhat smaller than the P. maculata wing, although this might be explained by sexual dimorphism. Its preserved venation is similar enough to that of the P. maculata holotype that it is likely closely related or perhaps even conspecific as the tentative identification on the label indicates. The basal branching of RA and RP is notably like that of the P. maculata type specimen. The posterior margin of the wing and CuPa��, CuPb, and 1A are not preserved, but the distal portions of the branches of CuA+CuPa�� appear long, at a low angle to the wing length. We further evaluated the angle of these veins in Zeuneroptera, whose sole fossil is missing its distal posterior portion. Sharov���s reconstruction (1962: Fig. 402, redrawn here as Fig. 2A) presumes a quite wide wing with the reconstructed portions of CuPa��, CuPb, and 1A straightening in the missing portion from their curves in their preserved portions. In this interpretation, these veins are long as in the Stenopelmatoidea. This missing portion could, however, have been narrower than Sharov speculated, and if these veins continue their curvature as in his drawing of their preserved portions (we have not seen the fossil or its modern photographs; access to collections at the Natural History Museum, London, is restricted during the COVID-19 pandemic), then they terminate more basally, not as in the Stenopelmatoidea (Fig, 2B). Both reconstructions are possible, and so the angles and termination points of these veins are unknown. In the partial tegmen of Albertoilus, CuA+CuPa��, CuPa�� and possibly CuPb (but possibly not 1A?) appear to be angled low to the margin and most likely long (Fig. 3C). Although there are three fossil tegmina known of the new genus Republicopteron, none have the distal portions of these four veins preserved. Their preserved portions suggest, however, that they are long. In the holotype SR 00- 04-06 part of the anterior margin is preserved with which to align the fossil, indicating that at least MA, MP, and CuA+CuPa�� are long, oriented at a low angle to the length of the wing. In the more complete tegmen of the new genus and species Ypopteron nicola, the termination points of these veins are more proximal than in the Stenopelmatoidea and are like those of many Prophalangopsidae: only two of five branches of CuA+CuPa�� end in the distal quarter of the wing, and CuPa�� ends mid-wing (Fig. 4). Note that the distal portions of the basal branch of CuA+CuPa��, CuPa��, CuPb, and the anal veins are upturned by a fold of the wing as in the P. maculata holotype. These four veins then might or might not conform with Grochov���s diagnosis of the Stenopelmatoidea in Palaeorehnia and Zeuneroptera, probably do in Albertoilus and Republicopteron, and do not in Ypopteron. By the strong similarity of all other aspects of the venation of these five genera listed in our emended diagnosis below, we group them as a taxon of unknown superfamily affinity, suspecting that these veins might have a range of lengths among them, perhaps varying from the hagloid-type to the stenopelmatoid-type. If this is so, it would be in concordance with Gorochov (1995), who considered the venation of Zeuneropterinae to be intermediate between the Stenopelmatoidea and Hagloidea. Like the Prophalangopsidae, CuA+CuPa�� has numerous branches in the Zeuneropterinae, further excluding it from the Stenopelmatoidea (including the Anostostomatidae), where there are no more than two (Gorochov 1995, 2001). With the restoration of Palaeorehnia to the taxon, its name reverts to Palaeorehniinae. As it is not then associated with any family, we raise it to the family level and treat it as the Palaeorehniidae Zeuner stat. nov., defining it by emending Gorochov���s diagnosis (1995, part 1, page 126) of the Zeuneropterinae as follows. Emended diagnosis. The tegmen of Palaeorehniidae have venation most like that of female Prophalangopsidae (e.g., CuPb and 2A rather equally bowed away from 1A basally), but may be distinguished from them most easily by the following. 1: CuPa�� (Fig. 3, red) not aligned with CuPa (Fig. 3, dark blue), angled toward M before its branching, toward anterior margin (Y. nicola: weakly; all others: more so) [all Hagloidea: CuPa�� aligned with CuPa, subparallel to M before M branching, directed toward posterior margin]; 2: basal branch of CuA+CuPa�� (Fig. 3A, light blue) not aligned with free CuA [Prophalangopsidae: almost always aligned, but see discussion]; 3: CuPa�� oblique to CuA+CuPa�� not aligned (Fig. 3, purple), these, with free CuA (Fig. 3, green) form a ���Y rotated counterclockwise [Prophalangopsidae: CuPa�� aligned with CuA+CuPa�� distal basal branch as in character state 2; these, with free CuA form an ���X���]; 4: as a consequence of 1 and 3, space posterior to M+CuA (Fig. 3, yellow), twice or more width of space posterior to M immediately distal to it (Fig. 3, orange) [Prophalangopsidae: similar width, but narrowing distal to branching of M (Fig. 3, light brown) in many]; Type and included genera. Type genus: Palaeorehnia; included genera: Zeuneroptera, Ypopteron, Albertoilus, and Republicopteron. Discussion. In Prophalangopsidae, the basal branch of CuA+CuPa�� (see character state 2 of the diagnosis) is shifted basally in Gorochov���s (1996, Fig. 5A) drawing of the tegmen of Karatailus micropterus Gorochov and appears shifted slightly distally in a specimen of a male Ashangopsis daohugouensis Lin et al. (see Gu et al. 2010, Fig. 9.2), but is aligned with the free CuA in the female specimen (Fig. 9.4). The basal branch of CuA+CuPa�� appears not aligned in Prophalangosis obscura (Walker), see Liu et al. (2009, Fig. 1, their CuA2), but we believe this to be due to creasing of the wing; in a specimen that we examined, this basal branch originates at the distal end of the free CuA as in the above diagnosis, although it appears at a slightly different angle from it, also because of creasing of the wing. Two genera thought at times to belong to the Zeuneropterinae have been transferred to the Tettigoniidae: Lithymnetes (Th��obald) 1937 (Oligocene, France, now Archepseudophylla Nel et al.) (Nel et al. 2008), and Eodecticus Pongr��cz (Miocene, Croatia) (Zeuner 1939 and see Gorochov 1995). Wang et al. (2019) transferred Hylophalangopsis chinensis Lin and Huang from the Prophalangopsidae to the taxon, however, while it agrees with character state 2 of our diagnosis (basal branch of CuA+CuPa�� not aligned with free CuA), it disagrees with it and agrees with Prophalangopsidae by character states 1, 3, and 4. Although this wing resembles those of Prophalangopsidae in these ways, MA, MP, CuA+CuPa��, CuPb, CuPa�� appear probably long as in Stenopelmatoidea, not Hagloidea. A thorough analysis and placement of it to higher taxon is outside of the scope of this paper, but we are confident that it is not a member of the Palaeorehniidae. In Prophalangopsidae, details of the branching of CuA+CuPa�� vary within species and even between the left and right wings of an individual (Gu et al. 2009, Wang et al. 2017). We presume this is also true in the Palaeorehniidae; therefore, we do not include these in Table 1 (as above, however, that this vein has more than two branches is informative). We suspect that perhaps two or more of the genera of Palaeorehniidae should be synonymized and some of the character states listed in Table 1 considered species-level differences. We refrain from doing so here as premature because except for Republicopteron, they are only known from single specimens (UCM-18757 is only tentatively associated with P. maculata) and all are fragmentary. 1: basal portion of 3A curves away from margin, toward 2A: A, yes; B, no; 2: space posterior to 2A about as wide as space posterior to 1A: A, yes; B, about half width; C, much narrower; 3: R branches basally, about same level as does M: A, yes; B, R branches notably distal to M; 4: maximum width of space posterior to CuA+CuPa�� about twice that posterior to CuPa��: A, yes; B, about the same. 5: 1A straight to about level of CuA and CuPa�� joining, then curved: A, yes; B, at least twice that distance; C, evenly curved before and after this level; 6: 3A ends on margin before or at level of CuA and CuPa�� joining: A, yes; B, beyond; 7: CuA+CuPa�� basal to first branch shorter than CuPa��: A, yes; B, as long or longer. *likely B (but most of space posterior to 2A not preserved); ���likely A (but region basal to level of CuA and CuPa�� joining mostly not preserved); ���most likely B (but distal-most portion of 1A compared here not preserved); �� most likely before this level; �� likely beyond this level. The three specimens assigned to R. douseae are consistent in the character states listed in table 1. Although the length of CuA+CuPa�� before its first branching varies to some degree, it is always long, distinct from that of all other Palaeorehniidae. All three wings have dark membranes. The holotype has some spots, but much fewer than those of P. maculata. The two paratypes do not bear such spots, but this might be an artefact; compare the part and counterpart of P. maculata (Fig. 1A and 1B). The photograph and drawing of the tegmen of A. cervirufi by Kevan and Wighton (1983) are not very informative as to colouration, but they described the wing as bearing reasonably well developed maculations that are not definite in shape, although this pattern could also be an artefact. In Zeuner���s description of Z. scotica (as P. scotica), he makes no mention of colouration, and we know of no description or illustration of this. The wing of Y. nicola appears infuscate throughout, although preservation of the membrane varies across it., Published as part of Archibald, S. Bruce, Gu, Jun-Jie & Mathewes, Rolf W., 2022, The Palaeorehniidae (Orthoptera, Ensifera, " Zeuneropterinae "), and new taxa from the early Eocene Okanagan Highlands, western North America, pp. 559-572 in Zootaxa 5100 (4) on pages 560-565, DOI: 10.11646/zootaxa.5100.4.6, http://zenodo.org/record/6225002, {"references":["Cockerell, T. D. A. (1908) A fossil Orthopterous insect with the media and cubitus fusing. Entomological News, 19, 126 - 128.","Zeuner, F. E. (1937) Descriptions of new genera and species of fossil Saltatoria (Orthoptera). Proceedings of the Entomological Society of London, Series B, 6, 154 - 159 https: // doi. org / 10.1111 / j. 1365 - 3113.1937. tb 00314. x","Scudder, S. H. (1890) The Tertiary Insects of North America. Report of the United States Geological Survey of the Territories, 13, 1 - 734. https: // doi. org / 10.5962 / bhl. title. 2507","Cockerell, T. D. A. (1909) Description of Tertiary insects, VII. American Journal of Sciences, Series 4, 28, 283 - 286. https: // doi. org / 10.2475 / ajs. s 4 - 28.165.283","Kevan, D. K. M. & Wighton, D. C. (1983) Further observations on North American Tertiary orthopteroids from south-central Alberta, Canada. Canadian Journal of Earth Sciences, 20, 217 - 224. https: // doi. org / 10.1139 / e 83 - 019","Sharov, A. G. (1962) Orthoptera (Saltatoria). In B. B. Rohdendorf ed. Osnovy Paleontologii: Spravochnik dlya paleontologov i geologov SSSR, 9, Chlenistonogie, Tracheiny i Khelitserovye [Fundamentals of paleontology: reference book for paleontologists and geologists of the U. S. S. R. 9, arthropods, tracheates and chelicerates]. Izdatel'stvo Akademiya Nauk SSSR, Moscow, pp 146 - 157. [English translation 1971 published by the Smithsonian Institution, Washington, D. C., pp. 192 - 209]","Ragge, D. R. (1955) The wing-venation of the Orthoptera Saltatoria, with notes on Dictyopteran wing-venation. British Museum (Natural History), London, vi + 159 pp.","Kevan, D. K. McE. & Wighton, D. C. (1981) Paleocene orthopteroids from south-central Alberta, Canada. Canadian Journal of Earth Sciences, 18, 1824 - 1837. https: // doi. org / 10.1139 / e 81 - 170","Gorochov, A. V. (2001) the higher classification, phylogeny and evolution of the Superfamily Stenopelmatoidea. In: Field, L. H. (Ed.), The Biology of Wetas, King Crickets, and their Allies. CABI publishing, New York, pp. 1 - 540. https: // doi. org / 10.1079 / 9780851994086.0003","Bethoux, O. (2012) King crickets, raspy crickets and weta, their wings, their fossil relatives. Journal of Orthoptera Research, 21, 179 - 225. https: // doi. org / 10.1665 / 034.021.0206","Zeuner, F. E. (1939) Fossil Orthoptera Ensifera. British Museum (Natural History), London, 321 pp.","Gu, J. - J., Qiao, G. - X. & Dong, R. (2010) Revision and new taxa of fossil Prophalangopsidae (Orthoptera: Ensifera). Journal of Orthoptera Research, 19, 41 - 56. https: // doi. org / 10.1665 / 034.019.0110","Liu, X., Zhou, M., Bi, W. & Tang, L. (2009) New data on taxonomy of recent Prophalangopsidae (Orthoptera: Hagloidea). Zootaxa, 2026 (1), 53 - 62. https: // doi. org / 10.11646 / zootaxa. 2026.1.6","Nel, A., Prokop, J. & Ross, A. J. (2008) New genus of leaf-mimicking katydids (Orthoptera: Tettigoniidae) from the Late Eocene - Early Oligocene of France and England. Comptes Rendus Palevol, 7, 211 - 216. https: // doi. org / 10.1016 / j. crpv. 2008.02.008","Wang, H., Fang, Y., Li, S., Hou, X., Wang, B. & Zhang, H. (2019) Revisiting of the Paleocene orthopteran insect Hylophalangopsis chinensis Lin and Huang, 2006 in northern Tibet. Journal of Asian Earth Sciences, 175, 93 - 98. https: // doi. org / 10.1016 / j. jseaes. 2018.06.003","Gu, J., Zhao, Y. - Y. & Ren, D. (2009) New fossil Prophalangopsidae (Orthoptera, Hagloidea) from the Middle Jurassic of Inner Mongolia, China. Zootaxa, 2004 (1), 16 - 24. https: // doi. org / 10.11646 / zootaxa. 2004.1.2","Wang, H., Fang, Y., Zhang, Q., Lei, X., Wang, B., Jarzembowski, E. A. & Zhang, H. (2017) New material of Sigmaboilus (Insecta, Orthoptera, Prophalangopsidae) from the Jurassic Daohugou Beds, Inner Mongolia, China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 107, 177 - 183. https: // doi. org / 10.1017 / S 1755691017000172"]}

Ypopteron nicola new species (Fig. 4) Haglidae, Prophalangopsinae sp.: Archibald and Mathewes 2000: 1443, Fig. 3. Diagnosis. As for genus. Description. Holotype tegmen. As in diagnosis and: about 6.5 cm long as preserved (small portion of basalmost region missing); membrane apparently infuscate throughout; RA, RP branch distal branching of MA, MP; RP branching eight times, one branch branched again near margin; MA, MP branch at level of CuA joining CuPa��, distally parallel with branches of RP; CuA+CuPa�� with five branches preserved, probably total; no stridulatory file detected on CuPb; simple crossveins throughout proximal posterior portion; zigzag intercalated crossveins forming pentagons in apical portion of overlapping wing between branches of RP. Type specimen. Holotype. Q-1010, part (no counterpart), two or more wings, two almost perfectly overlapping. The anterior regions (RA, stem of RP, ScA, SCP and precostal region) are missing or covered by portions of one or more wing(s); labelled Ypopteron nicola Archibald, Gu, and Mathewes, holotype; collected by Jurgen Mathewes, 1989; deposited in the collections of Simon Fraser University. Locality and age. Quilchena, British Columbia; Coldwater Beds; mid Ypresian, 51.5 �� 0.4 Ma. old. Etymology. The specific epithet refers to the Nicola Valley, where the Quilchena locality is located., Published as part of Archibald, S. Bruce, Gu, Jun-Jie & Mathewes, Rolf W., 2022, The Palaeorehniidae (Orthoptera, Ensifera, " Zeuneropterinae "), and new taxa from the early Eocene Okanagan Highlands, western North America, pp. 559-572 in Zootaxa 5100 (4) on pages 566-567, DOI: 10.11646/zootaxa.5100.4.6, http://zenodo.org/record/6225002, {"references":["Archibald, S. B. & Mathewes, R. W. (2000) Early Eocene insects from Quilchena, British Columbia, and their paleoclimatic implications. Canadian Journal of Zoology, 78, 1441 - 1462. https: // doi. org / 10.1139 / z 00 - 070"]}

Osmylidia glastrai sp. nov. Fig. 1 Type material. Holotype SR 02-22-15 (part) collected by Joshua Glastra, August 24, 2002; deposited in SR. A wellpreserved almost complete forewing. Type locality and horizon. Exposure B 4131 of the Tom Thumb Member of the Klondike Mountain Formation in the town of Republic, Washington, U.S.A.; Eocene (late Ypresian). Etymology. From the surname of Joshua Glastra, collector of this fossil. Diagnosis. Forewings may be distinguished from those of other species of the genus by faint colour patterning (O. picta: strongly spotted; O. requieta: without spots). Description. Forewing ca. 17 mm long, ca. 5.7 mm wide. Costal space broad, with subcostal veinlets simple, closely spaced. Sc fused with RA far from wing apex. Sc+RA veinlets simple, closely spaced. Subcostal space narrow; crossveins not detected. RA space relatively narrow, with five preserved crossveins, rather regularly spaced; one distal crossvein probably not preserved. RP originates rather close to wing base, with 14 pectinate branches, one of these (RP10) deeply forked. M forked distad origin of RP1, proximad origin of RP2. All crossveins (beside one proximal) in radial and radio-medial spaces arranged in three gradate series: proximal series with five crossveins; intermediate series with eight crossveins; outer series with seven preserved crossveins (partly fragmentarily preserved). Basal crossvein in radio-medial space connects R, M. Both MA, MP pectinately branched distally, with three branches. Four crossveins between MA, MP; five preserved crossveins between MP, CuA. CuA pectinately branched, with three branches. CuP pectinately branched, with five preserved branches (probably six or seven in life). One preserved crossvein detected between CuA, CuP. One distal crossvein between CuP, A1. A1 relatively long, with five pectinate, simple branches. A2 relatively long, with four preserved pectinate, simple branches. Wing colouration: irregular spots throughout wing (surrounding some crossveins, in subcostal space and along margin), most prominently three between MP, CuA, one along margin near termination of CuP, A1, and two on pterostigma. Remarks. The species has a five-branched CuP and, therefore, is assigned to Osmylidia (see genus diagnosis) but only tentatively, as this character is unknown in one fossil genus, and Osmylidia is barely distinguished from extant genera solely by forewing venation., Published as part of Makarkin, Vladimir N., Archibald, S. Bruce & Mathewes, Rolf W., 2021, New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks, pp. 142-156 in Zootaxa 4980 (1) on pages 144-145, DOI: 10.11646/zootaxa.4980.1.9, http://zenodo.org/record/4883181

Osmylidia sp. Fig. 4 Material. Specimen RBCM P1567, collected by SBA July 23, 2001; deposited in RBCM. A poorly preserved incomplete hind wing on a piece of shale with a heteropteran (Hemiptera), a syrphid (Diptera), and two nematoceran pupae (Diptera). Locality and horizon. Ootsa Group shale exposed at Driftwood Canyon Provincial Park, British Columbia, Canada; Eocene (mid-Ypresian). Description. Hind wing 10.9 mm long as preserved (estimated complete length about 11.5 mm), ca. 5 mm wide (actual width probably less, wing deformed). Costal space relatively broad, with simple, relatively closely spaced subcostal veinlets. Sc fused with RA far from wing apex. Subcostal space incompletely preserved, narrow distally; crossveins not detected. RA space relatively narrow for most length, basally broadened, with three detected crossveins in proximal half. RP with 11 branches, all probably shallowly forked (their deep forks absent, distal portions not preserved). Fork of M not detected. MA probably deeply forked, MP probably pectinate (both fragmentarily preserved). Nearly all preserved crossveins in radial, radio-medial spaces arranged in intermediate, outer gradate series (both series incomplete) or single crossvein in position of proximal gradate series in other Protosmylinae; one crossvein in intermediate series adventitiously forked. Two preserved crossveins between posterior branch of MA, MP. CuA strongly pectinately branched, with eight branches. CuP relatively short, forked twice distally. Remarks. Although it differs from other species by its arrangement of crossveins between RA and RA, this incomplete specimen lacks many potentially diagnostic characters (e.g., the configuration of M), and we, therefore, do not name a species based on it., Published as part of Makarkin, Vladimir N., Archibald, S. Bruce & Mathewes, Rolf W., 2021, New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks, pp. 142-156 in Zootaxa 4980 (1) on page 149, DOI: 10.11646/zootaxa.4980.1.9, http://zenodo.org/record/4883181

Genus Osmylidia Cockerell, 1908 Osmylidia Cockerell, 1908: 342. Oligosmylus Kr��ger, 1913: 34. Protosmylus Kr��ger, 1913: 30, syn. nov. Type species. Osmylus requietus Scudder, 1890, by monotypy. Included species. Osmylidia donnae sp. nov., (mid Ypresian, Quilchena), O. glastrai sp. nov. (late Ypresian, Republic), O. taliae sp. nov. (late Ypresian, Green River Formation); Osmylidia sp. (mid Ypresian, Driftwood Canyon), O. requieta (late Priabonian, Florissant), O. picta (Hagen in Pictet-Baraban & Hagen, 1856) (Priabonian, Baltic amber), comb. nov. Revised diagnosis. Separated from other genera of the subfamily by hind wing MA deeply forked [shallowly forked in other genera (unknown in Mesozoic Sogjuta, Jurosmylus and Protosmylina)], female procoxa with a large process (clearly seen only in O. picta) [absent in other genera]. Forewing venation differs from that of other fossil genera by greater number of CuP branches (5���7) [��� 4 in Sogjuta, Juraheterosmylus, Jurosmylus, Pseudosmylidia; unknown in Protosmylina]. Occurrence. Eocene (Ypresian to Priabonian) of North America and northern Europe. Remarks. The forewing character states of extant genera vary greatly among their species, covering almost the entire range of those of the family, but the shallowly forked hind wing MA is consistent in these genera. The proximal crossveins in the forewing radial space of Osmylidia requieta are arranged rather irregularly (see e.g., Carpenter 1943: Fig. 2). This is associated with at least eight and as many as ten crossveins between RA and RP while all other species of the genus possess at most six. We interpret this greater number of crossveins in these spaces and their arrangement as a species character., Published as part of Makarkin, Vladimir N., Archibald, S. Bruce & Mathewes, Rolf W., 2021, New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks, pp. 142-156 in Zootaxa 4980 (1) on page 144, DOI: 10.11646/zootaxa.4980.1.9, http://zenodo.org/record/4883181, {"references":["Cockerell, T. D. A. (1908) Fossil Osmylidae (Neuroptera) in America. The Canadian Entomologist, 40, 341 - 342. https: // doi. org / 10.4039 / Ent 40341 - 10","Kruger, L. (1913) Osmylidae. Beitrage zu einer Monographie der Neuropteren-Familie der Osmyliden. II. Charakteristik der Familie, Unterfamilien und Gattungen auf Grund des Geaders. Stettiner Entomologische Zeitung, 74, 3 - 123.","Scudder, S. H. (1890) The Tertiary Insects of North America. Report of the United States Geological Survey of the Territories, 13, 1 - 734. https: // doi. org / 10.5962 / bhl. title. 2507","Pictet-Baraban, F. J. & Hagen, H. A. (1856) Die im Bernstein befindlichen Neuropteren der Vorvelt. In: Berendt, C. G. (Ed.), Die im Bernstein befindlichen organischen Reste der Vorwelt gesammelt, in verbindung mit mehreren bearbeitet und herausgegeben von Dr. Georg Carl Berendt. Band 2. Abteilung 2. Nicholaischen Buchhandlung, Berlin, pp. 41 - 125.","Carpenter, F. M. (1943) Osmylidae of the Florissant shales, Colorado (Insecta-Neuroptera). American Journal of Science, 241, 753 - 760. https: // doi. org / 10.2475 / ajs. 241.12.753"]}

Osmylidia donnae sp. nov. Fig. 2 Type material. Holotype Q-0317, collected by RWM, October 2001; deposited in SFU. A rather well-preserved almost complete hind wing. Type locality and horizon. Quilchena, British Columbia, Canada; Eocene (mid-Ypresian). Etymology. From the first name of Donna Mathewes, for her support of fieldwork at Quilchena. Diagnosis. May be distinguished from hind wings of other species of the genus by long distal crossvein between posterior branch of MA, MP [other species: this crossvein short or lacking as these veins fused for short distance], and membrane slightly infuscate [other species: hyaline]. Description. Hind wing 13.5 mm as preserved (estimated complete length about 14 mm), 4.3 mm wide. Costal space relatively narrow, with simple, relatively closely spaced subcostal veinlets. Sc fused with RA far from wing apex. Sc+RA veinlets simple, closely spaced. Subcostal space narrow; crossveins not detected. RA space relatively narrow, with four widely spaced crossveins. RP with 10 branches, RP8 deeply forked, other branches shallowly forked. Basal crossvein 1r-m preserved fragmentarily, near RP. M forked near base wing, slightly proximad origin of RP. MA deeply forked, proximad level of proximal crossvein between RP1, RP2. All crossveins in radial, radiomedial spaces arranged in three gradate series: proximal series with two crossveins; intermediate series with five crossveins; outer series with eight preserved crossveins (region partly missing or damaged). Anterior branch of MA pectinately branched with four branches; posterior branch of MA with terminal fork, one incompletely preserved branch. MP pectinately branched with two branches. Two crossveins (one fragmentarily preserved) between MA, MP proximad forking of MA; three crossveins between anterior, posterior branches of MA, between posterior branch of MA, MP (including long distal crossvein, i.e., posterior branch of MA, MP not fused). CuA strongly pectinately branched, with seven preserved branches (probably eight in life). CuP relatively short, forked once distally. Two crossveins between CuA, CuP. 1A with two pectinate branches. Membrane mostly slightly infuscate (as preserved; perhaps uniformly infuscate in life?)., Published as part of Makarkin, Vladimir N., Archibald, S. Bruce & Mathewes, Rolf W., 2021, New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks, pp. 142-156 in Zootaxa 4980 (1) on page 146, DOI: 10.11646/zootaxa.4980.1.9, http://zenodo.org/record/4883181

Osmylidia taliae sp. nov. Fig. 3 Type material. Holotype UCM 67996, collected by David Kohls, 4 March 2004; deposited in UCM. A poorly preserved incomplete specimen with at least small portions of all four wings. Type locality and horizon. UCM locality 2005025 (Anvil Points Kohls site); Garfield County, Colorado, U.S.A. Parachute Creek Member of the Green River Formation; late Ypresian (early Eocene). Etymology. From the first name of Talia Karim, for her long-standing helpful support of paleoentomology. Diagnosis. May be distinguished from other species of the genus by forewing stem of RP distally pectinately branched [other species: dichotomously branched]; MP not branched at outer gradate series [other species: branched]; hind wing posterior branch of MA fused with MP for short distance [other species (except some specimens of O. picta): separate, connected by crossvein]. Description. Body poorly preserved, ca. 9 mm long; details not discernable. Forewing ca. 14 mm long, ca. 4.8���5.0 mm wide. Costal space broad, with subcostal veinlets simple, closely spaced. Sc fused with RA far from wing apex. Subcostal space narrow in distal portion; crossveins not detected. RA space relatively broad medially, with two preserved crossveins. RP with 11 pectinate branches, all shallowly forked. All preserved crossveins in radial to medio-cubital spaces arranged in three gradate series: proximal series with three preserved crossveins; intermediate series with eight preserved crossveins; outer series with 11 preserved crossveins (probably 14 in life). MA pectinately branched distally, probably with three branches. MP probably shallowly branched (apex not preserved), at least not branched at outer gradate series or proximad it. CuA pectinately branched, with three preserved branches. Wing colouration: faint spots at pterostigma and along apical margin; some crossveins in gradate series appear surrounded by fuscous colouration. Hind wing 7.5 mm long as preserved (estimated complete length ca. 13 mm), ca. 3 mm wide as preserved (estimated complete width ca. 3.5 mm). Costal space moderately broad, with simple, relatively closely spaced preserved subcostal veinlets. Three closely spaced crossveins preserved in RA space medially. RP with five partially preserved branches. MA deeply forked, distally pectinately forked, with at least three branches. MP pectinately forked with two branches. CuA fragmentarily preserved. Crossveins in radial, radio-medial spaces arranged in two preserved gradate series: intermediate series with four preserved crossveins; outer series with six preserved crossveins. Two preserved crossveins between posterior branch of MA, MP., Published as part of Makarkin, Vladimir N., Archibald, S. Bruce & Mathewes, Rolf W., 2021, New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks, pp. 142-156 in Zootaxa 4980 (1) on pages 147-148, DOI: 10.11646/zootaxa.4980.1.9, http://zenodo.org/record/4883181

Subfamily Protosmylinae Kr��ger, 1913 Type genus. Protosmylus Kr��ger, 1913 (= Osmylidia Cockerell, 1908). Emended diagnosis (based on wings). Wings may be distinguished from those of other subfamilies of Osmylidae by a combination of the following. Forewing: Subcostal veinlets almost always simple (very rarely some forked) [Osmylinae: mostly forked]; trichosors along almost entire wing margin [Gumillinae: restricted to apical region]; one subcostal crossvein, positioned basally [Porisminae: numerous along entire subcostal space]; all or nearly all crossveins in radial space forming three to four gradate series (except some Lysmus) [all or nearly all crossveins (Gumillinae, Porisminae, Eidoporisminae) or proximal crossveins (other subfamilies) irregularly arranged]; M forked at level or only slightly proximad termination of A1 [Osmylinae, Kempyninae: much distad; Stenosmylinae, Porisminae, Eidoporisminae: much proximad]; between CuA, CuP ��� 5 crossveins (up to 8 in some Lysmus) [Spilosmylinae: about 10 or more]; CuA pectinate, with 2���6 branches; CuP with 3���7 pectinate branches (up to 9 in some Lysmus) [Spilosmylinae: 8���15]. Hind wing: MP lacks anteriorly directed basal spur-like process [Spilosmylinae: present]; CuP relatively short, shallowly (not pectinately) branched or simple [Kempyninae, Osmylinae, Stenosmylinae, Porisminae, Eidoporisminae: long, pectinately branched]. Genera included. Ten genera: ��� Sogjuta Martynova, 1958 (Early to Late Jurassic, Kyrgyzstan and Mongolia), sit. nov.; ��� Juraheterosmylus Wang et al., 2010 (late Middle / early Late Jurassic, China); ��� Jurosmylus Makarkin & Archibald, 2005 (Late Jurassic, Kazakhstan); ��� Protosmylina Jepson et al. 2009 (Early Cretaceous, England); ��� Osmylidia (= Protosmylus, Priabonian, Baltic amber, syn. nov.), and ��� Pseudosmylidia Makarkin, 2017 (Priabonian, North America and Europe); Heterosmylus Kr��ger, 1913, Gryposmylus Kr��ger, 1913, and Lysmus Nav��s, 1911 (extant, all southeastern Asia); Paryphosmylus Kr��ger, 1913 (extant, Ecuador). Remarks. The venation of some species of Lysmus is very similar to that of the Spilosmylinae, e.g., L. harmandinus Nav��s, 1910 (see Makarkin 1985: Fig. 3; Winterton et al. 2019: Fig. 34). Moreover, this genus had long been considered a member of the Spilosmylinae. Lysmus, however, lacks an important synapomorphy of Spilosmylinae, the presence of an anteriorly directed basal spur-like process in the hind wing MP. Also, the male genitalia of Lysmus are typical for Protosmylinae (see Wang 2010). According to Article 40.1 of ICZN, a family-group name is not replaced with a new family-group name when the name of its type genus is considered to be a junior synonym of another genus. Hence, Protosmylinae retains this name following designation of Protosmylus as a junior synonym of Osmylidia., Published as part of Makarkin, Vladimir N., Archibald, S. Bruce & Mathewes, Rolf W., 2021, New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks, pp. 142-156 in Zootaxa 4980 (1) on pages 143-144, DOI: 10.11646/zootaxa.4980.1.9, http://zenodo.org/record/4883181, {"references":["Kruger, L. (1913) Osmylidae. Beitrage zu einer Monographie der Neuropteren-Familie der Osmyliden. II. Charakteristik der Familie, Unterfamilien und Gattungen auf Grund des Geaders. Stettiner Entomologische Zeitung, 74, 3 - 123.","Cockerell, T. D. A. (1908) Fossil Osmylidae (Neuroptera) in America. The Canadian Entomologist, 40, 341 - 342. https: // doi. org / 10.4039 / Ent 40341 - 10","Martynova, O. M. (1958) New insects from the Permian and Mesozoic deposits of the USSR. In: Sharov, A. G. (Ed.), Materialy k Osnovam Paleontologii [Materials to \" Fundamentals of Paleontology \"]. Vol. 2. Academy of Sciences of the USSR, Moscow, pp. 69 - 94. [in Russian]","Wang, Y. J., Liu, Z. Q., Ren, D. & Shih, C. K. (2010) A new genus of Protosmylinae from the Middle Jurassic of China (Neuroptera: Osmylidae). Zootaxa, 2480 (1), 45 - 53. https: // doi. org / 10.11646 / zootaxa. 2480.1.4","Makarkin, V. N. & Archibald, S. B. (2005) Substitute names for three genera of fossil Neuroptera, with taxonomic notes. Zootaxa, 1054 (1), 15 - 23. https: // doi. org / 10.11646 / zootaxa. 1054.1.2","Jepson, J. E., Makarkin, V. N. & Jarzembowski, E. A. (2009) New lacewings (Insecta: Neuroptera) from the Lower Cretaceous Wealden Supergroup of southern England. Cretaceous Research, 30, 1325 - 1338. https: // doi. org / 10.1016 / j. cretres. 2009.07.012","Makarkin, V. N. (2017) A remarkable new genus of Protosmylinae (Neuroptera: Osmylidae) from late Eocene Florissant, Colorado. Zootaxa, 4268 (4), 581 - 587. https: // doi. org / 10.11646 / zootaxa. 4268.4.9","Navas, L. (1911) Nevropteres nouveaux de l'extreme Orient. Russkoe Entomologicheskoe Obozrenie [Revue Russe d'Entomologie], 11, 111 - 117.","Makarkin, V. N. (1985) Review of the family Osmylidae (Neuroptera) of the fauna of the USSR. In: Lehr, P. A. & Storozhenko, S. Yu. (Chief Eds.), Taxonomy and ecology of arthropods of the Far East of the USSR. 129. Abstract. Far Eastern Scientific Centre of the Academy of Sciences of the USSR, Vladivostok, pp. 35 - 47. [in Russian]","Winterton, S. L., Martins, C. C., Makarkin, V. N., Ardila-Camacho, A. & Wang, Y. J. (2019) Lance lacewings of the world (Neuroptera: Archeosmylidae, Osmylidae & Saucrosmylidae): a review of living and fossil genera. Zootaxa, 4581 (1), 1 - 99. https: // doi. org / 10.11646 / zootaxa. 4581.1.1"]}

Okanagrion threadgillae Archibald & Cannings, new species Figs. 19–20. Diagnosis. Wings separated from those of O. beardi, O. dorrellae by MP, CuA sub-parallel at terminus on posterior margin [O. beardi: distinctly wider, O. dorrellae: widening, but less distinctly as CuA zigzagged distally (O. angustum, not known); from O. dorrellae by MP linear to margin [O. dorrellae: zigzagged near margin]; from O. angus- tum, O. beardi by IR2 origin between RP1-2, RP3-4 [O. angustum, O. beardi: origin on RP1-2]; from O. dorrellae by wider wing: length (arculus to apex)/ maximum width, 3.1 [O. dorrellae: 3.7], nodus to apex 2.5 times maximum width [although the wing of O. angustum is less complete, it is distinctly narrower, compare angle of posterior margin to anterior margin preserved between levels of nodus, origin of RP2]; width at level of two cells proximal to origin of RP 2 in O. angustum 75% that of O. threadgillae; from O. liquetoalatum by wing shape: while length of arculus to nodus is equivalent, arculus to apex is about 2 mm longer in O. threadgillae; from O. liquetoalatum, O. dorrellae by wing shape: length (arculus to apex) / maximum width 3.1 [O. liquetoalatum: 2.6; O. dorrellae 3.7; others similar to O. threadgillae (O. lochmum, O. worleyae, O. angustum not comparable by preservation)]; wing length (arculus to apex) / maximum width, O. dorrellae 3.7; from O. hobani by main veins less deeply curved near margin; from O. worleyae further by shape: posterior margin more deeply curved; from O. lochmum by RA–RP1 space becomes two cells wide about two cells proximal to pterostigma [O. lochmum over seven cells proximal]. Further separated by colouration (sex unknown): proximal approx. 2/5 hyaline, dark apical to this except narrow lighter fascia just distal to mid-way between nodus, pterostigma, extending from anterior margin to mid-wing; size and extent of light fascia differs from: O. hobani about three times as wide, extending to or almost to posterior margin; O. angustum apparently similar to O. hobani as preserved; O. beardi, O. lochmum, wider than in O. threadgillae by small but definite amount, extending closer to posterior margin; O. dorrellae similar width as O. beardi, O. lochmum, probably extending to mid-wing (but damage in this region); O. liquetoalatum: hyaline throughout; O. worleyae: lightly infuscate throughout known wing. Type material. Holotype: SR 98-12-10 (Republic A0307, collected by Carolyn Threadgill 8.iv.1998): two well preserved, complete wings; disarticulated, in close association; with colour patterning clearly preserved; housed in the Stonerose Interpretive Center collection. Description. Holotype, SR 98-12-10, forewing (Figs. 19A, 20A): distal to nodus rather wide-oval, symmetrical around longitudinal line from apex; posterior margin slightly widens about level of quadrangle; 30.5 mm long, 27.6 mm arculus to apex, 22.5 mm nodus to apex; 18.0 mm origin of RP2 to apex, 21.5 mm arculus to base of pterostigma, 16.5 mm nodus to base of pterostigma; 9.0 mm wide. Nodus at 28% wing length. Colouration as in diagnosis. Pterostigma anterior, posterior margins oblique; length 3.5 times width; subtends 5.5 cells; no oblique brace vein. 40 crossveins in postnodal space (complete), the distal-most one joining pterostigma (hyperstigmal crossvein); not aligned with postsubnodal crossveins. Maximum three cells wide in much of costal space distal to pterostigma. RA meets margin at apex; RA–RP1 space narrows near margin. RP1–IR1 space goes to two cells wide 10 cells distal to origin of IR1. Cells in apical portion increasingly dense, many small cells along margin. IR1 origin nine cells distal to origin of RP2; IR1–RP2 space becomes two cells wide about ten cells distal to origin. RP2 origin 6.5 cells distal to origin of IR2. No crossvein O. IR2 origin at subnodus. R3-4 origin less than 2/3 distance between arculus, nodus. Arculus immediately proximal to Ax2. Ax0 not detected. All main veins linear except proximal portion of IR1. MP, CuA slightly divergent at terminus on posterior margin. CuA ends at level about half way between nodus, pterostigma; CuA–A space maximum five cells wide. Hind wing (Figs. 19B, 20B): As for forewing, except: Posterior margin evenly curved about level of quadrangle, does not slightly widen; 26.5 mm arculus to apex, 21.7 mm nodus to apex, 17.7 mm origin of RP2 to apex, 20.6 mm arculus to base of pterostigma, 15.7 mm nodus to base of pterostigma; 8.5 mm wide. Nodus at 30% wing length. Pterostigma length 3.5 times width; subtends six cells. 35 crossveins preserved (presume 36 total) in postnodal space, distal-most three hyperstigmal. One column four cells wide, mostly three in costal space distal to pterostigma. RP1–IR1 space goes to two cells wide 11 cells distal to origin of IR1. IR1 origin six cells distal to origin of RP2; IR1–RP2 space becomes two cells wide eleven cells distal to origin. RP2 origin probably five cells distal to origin of IR2. Origin of RP3-4 slightly proximal to position in forewing. MP, CuA sub-parallel at terminus on posterior margin. CuA–A space maximum six cells wide. Etymology. The specific epithet is a patronymic formed from the surname of Carolyn Threadgill, the collector of the holotype, recognising her contribution. Range and age. Republic locality A0307 of the Tom Thumb Tuff Member of the Klondike Mountain Formation; latest Ypresian. Discussion. We presume the top wing of Figs. 19A and 20A to be a forewing and the bottom wing of these (Figs. 19B and 20B) to be a hind wing, as discussed above., Published as part of Archibald, Bruce, Cannings, Robert A., Erickson, Robert J., Bybee, Seth M. & Mathewes, Rolf W., 2021, The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America, pp. 1-133 in Zootaxa 4934 (1) on pages 43-44, DOI: 10.11646/zootaxa.4934.1.1, http://zenodo.org/record/4558796

Reassessment of the Dysagrionidae Taxon concepts based on the presence of accessory antenodal crossveins. The Dysagrioninae was originally proposed by Cockerell (1908a) as a subfamily of Agrionidae (Zygoptera), comprised of the two extinct genera Dysagrion and Phenacolestes. He distinguished it from other agrionids solely by the presence of accessory antenodal crossveins, i.e., in the costal space between Ax2 and the nodus: two in Dysagrion and three in Phenacolestes (P. mirandus Cockerell from the Priabonian shale of Florissant, Colorado was then its only confirmed species, Fig. 2). Calvert (1913) examined six specimens of P. mirandus, five of which had the antenodal region preserved, and found that three of these had three accessory antenodal crossveins, and the other two had two. He agreed with Campion (1913) that this variability made Cockerell���s Dysagrioninae untenable. Campion thought that P. mirandus most closely resembles the extant Neotropical Thaumatoneura McLachlan, which also has a variable number of accessory antenodal crossveins. Tillyard & Fraser (1939), Fraser (1957) and Nel & Paicheler (1994) did not recognise Cockerell���s Dysagrioninae, placing Phenacolestes in the Amphipterygidae (Tillyard & Fraser 1939) and Pseudolestidae (Thaumatoneurinae) (Fraser 1957; Nel & Paicheler 1994). Modern photographs clearly confirm the presence of three accessory antenodal crossveins in the holotype of P. mirandus (US National Parks Service website, see our drawing from this photograph, Fig. 2). Cockerell (1908a) also described a new species from Florissant as Phenacolestes ? parallelus Cockerell, a tentative member of the genus as the region of the diagnostic accessory antenodal crossveins is not preserved (University of Colorado: UCM 4503, part and counterpart). Subsequently discovered specimens confirmed that the species is a Phenacolestes: a proximal half of a wing (Yale Peabody Museum: YPM IP 220974) has four accessory antenodal crossveins (Cockerell 1908b) and a third, almost complete specimen (University of Colorado: UCM 4545), also from Florissant (Cockerell 1908c), has three in both forewings and three in both hind wings. Here, we figure these (Figs. 2, 3) as, to our knowledge, no specimen of the species has been previously illustrated other than by a poorquality photograph of UCM 4545 (Cockerell 1908c, his Fig. 4). The third species attributed to the genus, P. ? coloratus (Hagen) (mid-Miocene, Radoboj, Croatia) is poorly known, inadequately described, and we know of no published drawing or photograph (Hagen 1848; and see Nel & Paicheler 1994; Nel et al. 2005b). We treat it as a possible member of the genus (adding the question mark), family, and suborder pending examination of the specimen. The discussions of the above authors concerned the appropriateness of basing the taxon on a variable character state, i.e., two or three accessory antenodals in P. mirandus (the variability in P. parallelus described above was not known to them); the presence of two such crossveins in Dysagrion, however, was not doubted. Although Scudder���s original drawings should have established that this is not always the case, it remained unquestioned from the time of Cockerell���s works cited above to the present day. Scudder���s drawing of Dysagrion fredericii Scudder (Ypresian, Green River Formation, Wyoming) (1890: plate 6, Fig. 9) type specimen 4167/4168 (now MCZ numbers 381A, B) shows two accessory antenodal crossveins; however, there are none in his drawing (1890: plate 6, Fig. 3) of the type of D. packardii Scudder (Green River Formation, Wyoming) (Scudder cites the specimen as Packard number 146), a copy of which had already appeared in a work by Zittel (1885: Fig. 979) which Scudder edited. Tillyard & Fraser (1939: Fig. 3.1) provided a drawing of the wing (re-drawn from Scudder?), adding two accessory antenodal crossveins, which Fraser subsequently reproduced (1957: Fig. 34.1) (the specific epithet spelled ��� packardi ��� in both). Although Carpenter had the Museum of Comparative Zoology (MCZ) specimens available to him, his drawing of D. packardii (1992, his Fig. 52.1) included these presumptive accessory antenodal crossveins, by which he in part distinguished the genus. Bechly (1996) based his tribe Congqingiini (Congqingia Zhang and Petrolestes Cockerell) on its members lacking accessory antenodal crossveins, in contrast to his Dysagrionini (Dysagrion and Phenacolestes), reported to possess them as above. To resolve confusion as to the presence, and so the diagnostic usefulness, of accessory antenodal crossveins in Dysagrion species, considering the prominent role they have played in understanding their relationships at various taxonomic levels, we examined new, high resolution photographs of Scudder���s original Dysagrion specimens housed in the collections of the MCZ. The location of the holotype of D. packardii (Fig. 4C, re-drawn here from Scudder 1890, plate 6 Fig. 3) is unknown, but we were able to examine new photographs of cotype MCZ 4656 (Packard number 252) (Fig. 4A, B). In this specimen, the proximal portion of the antenodal region is missing, but the region where these crossveins should occur, if present, is reasonably well preserved. We did not detect any (either dry or wetted with ethanol), consistent with Scudder���s original description and contra subsequent authors. We also examined new photographs of the D. lakesii Scudder (Green River Formation, Wyoming) type specimen MCZ 4101 (Packard number 259, not previously illustrated) (Fig. 5); these also show no evidence of such accessory crossveins, which should be readily detectible if present. In the type of D. fredericii (Scudder���s number 4167/4168, now MCZ numbers 381A, B), we see one weakly preserved accessory crossvein, which we indicate by a dotted line in Fig. 6, but we doubt there are two, differing from Scudder���s drawing (1890) as discussed by Cockerell (1908a). Tillyard & Fraser (1939) and Fraser (1957: Fig. 34.2, spelled ��� frederici ���) subsequently reproduced Scudder���s drawing of this specimen with two accessory crossveins. We also examined photographs of MCZ 382, 1497, and 4147, but these fossils are fragmentary and do not possess this region. Cockerell and subsequent authors (as above) were then incorrect in asserting that the presence of accessory antenodal crossveins characterises Dysagrion and justifies its association with Phenacolestes. Seven genera have been subsequently associated with Dysagrion and Phenacolestes as forming the current Dysagrionidae: Primorilestes, Petrolestes, Congqingia, Electrophenacolestes Nel & Arillo, Burmadysagrion Zheng et al., Electrodysagrion, and Palaeodysagrion (we exclude the latter three, see below) (Figs. 7���9). None of these except Electrophenacolestes possess an accessory antenodal crossvein. None of the new species that we associate here with the family (below) possess them. A presence of accessory crossveins is not, therefore, a useful character state on which to base this family concept, define constituent subfamilies, nor characterise the genus Dysagrion. Subsequent definitions of Dysagrionidae. Some or all of the above genera and sometimes Thaumatoneura, Eodysagrion Rust et al., and others have been grouped in various family configurations, including in the Agrionidae, Amphipterygidae, Pseudolestidae, Megapodagrionidae, as subfamilies of Thaumatoneuridae (Dysagrioninae and Thaumatoneurinae: e.g., Bechly 1996; Nel et al. 2005 a, 2005b; Nel & Arillo 2006), or of the Dysagrionidae (Dysagrioninae, Thaumatoneurinae and Eodysagrioninae: Rust et al. 2008; Dysagrioninae, Burmadysagrioninae and Eodysagrioninae: Zheng et al. 2016a, 2016b, 2017; and independent of Thaumatoneura and Eodysagrion: Garrouste & Nel 2015; Nel et al. 2016; Huang et al. 2017). Bechly (1996) distinguished the taxon as Dysagrioninae (within Thaumatoneuridae: Dysagrion, Petrolestes, Phenacolestes and Congqingia) by quadrangle shape as its single defining trait, although he did not describe this, and noted that it is also present in Sieblosiidae. Nel & Arillo (2006) agreed, further distinguishing it by the combination of: antesubnodal space without crossveins; absence of the oblique vein O; CuA���A space very broad; the base of IR2 positioned proximal to the nodus; RP3-4 arising between the arculus and nodus closer to nodus; and the base of RP2 being in a very distal position. Nel & Fleck (2014) closely followed this definition. Rust et al. (2008) erected the Eodysagrioninae for Eodysagrion as an equal rank taxon with Dysagrioninae and Thaumatoneurinae within the Dysagrionidae. The Dysagrionidae is currently not considered to be close to the Thaumatoneuridae. Garrouste & Nel (2015) questioned a close relationship between them because of inconsistencies in positions of the nodus and bases of branches of RP and IR2, the difference in shape of their quadrangles and of their leg spines and thoracic skewedness, and as an incomplete interpleural suture is present in Petrolestes, but it is complete in Thaumatoneura. They further suggested that Dysagrionidae might not even belong in the Zygoptera. Huang et al. (2017) explicitly separated them, elevating Dysagrioninae and Thaumatoneurinae to family level. They rejected their putative synapomorphies of costal margin curvature and lack of crossveins in the antesubnodal space as weakly supported and most likely convergent, also discussing their difference in quadrangle shape. They further considered Eodysagrion as a provisional member of the Thaumatoneuridae, among their reasons further stressing the importance of quadrangle shape, which is rectangular in Eodysagrion like that of Thaumatoneura, not Dysagrionidae. Subfamilies of Dysagrionidae. Huang et al. (2017), in elevating the Dysagrioninae to family status, also elevat- ed the former tribes Petrolestini and Dysagrionini to subfamilies: Petrolestinae, comprised of Petrolestes (P. hendersoni Cockerell: Ypresian Green River Formation, Colorado, United States of America; P. messelensis Garrouste & Nel: Lutetian, Grube Messel, Germany) and Congqingia (C. rhora, Aptian, i.e., Early Cretaceous, Laiyang Formation, Shandong, China; recent age estimate: Zhou et al. 2020) (Fig. 8); and their revised concept of Dysagrioninae, comprised of Dysagrion, Phenacolestes, Primorilestes (P. violetae Nel et al.: Priabonian, Bolshaya Svetlovodnaya (Biamo), Primorye, Russia; P. madseni Rust et al.: earliest Ypresian Fur Formation, Denmark; P. magnificus Nel et al.: Miocene, Satovcha Graben, Sivik Formation, Bulgaria), and Electrophenacolestes (E. serafini: Priabonian, Baltic amber, Poland) (Fig. 7). Bechly (1996) separated these (as tribes) by Petrolestinae having: 1, no accessory antenodal crossveins; 2, IR1 shortened and strongly zigzagged; and 3, the bases of RP3-4 and IR2 situated midway between nodus and arculus [Dysagrioninae: origin of RP 3-4 in middle third between arculus and nodus, usually at about two-thirds the distance; IR2 originates at or very near subnodus]. However, character state 1 is, as above, variable and not informative. Nel et al. (2005b) rejected character state 2, as this region is not preserved in the type species Petrolestes hendersoni and is not consistent in other specimens. Character state 3 is the sole trait separating these taxa. In the Dysagrioninae the origin of RP3-4 is in the middle third between the arculus and nodus, usually at about two-thirds of the distance, and IR2 originates at or very near the subnodus. In the Petrolestinae, the origin of RP3-4 is more proximally positioned, closer to or at the point midway between the arculus and subnodus, and IR2 originates close to or on RP3-4. There are no currently recognised taxa of tribe-level rank of the Dysagrionidae. Emended diagnosis of the Dysagrionidae. The wings of Dysagrionidae are easily separated from those of other Cephalozygoptera and tentative Cephalozygoptera (new suborder, see below) by a combination of: 1- oblique crossvein O absent [Sieblosiidae: present]; 2- arculus at or immediately proximal to Ax2 [Whetwhetaksidae: nearer to Ax1]. They are distinguished from similar appearing Zygoptera by a combination of the above and the following character states 3���9 of the wings, slightly amended from the diagnoses of Nel & Arillo (2006) and Nel & Fleck (2014), and 10, of the head: 3- quadrangle broad, distal side longer than proximal side, posterior side longer than anterior, distal-posterior angle oblique, proximal-anterior angle usually about 90��; 4- nodus positioned at least a quarter wing length, usually more; 5- anterior anal vein separates from posterior anal vein briefly before joining CuP (i. e., is briefly free distal to petiole); 6- RP3-4 originating roughly between one third to two thirds length from arculus to subnodus, usually about two thirds (Primorilestes violetae furthest, just distal to middle third, but P. madseni within middle third); 7- antesubnodal space without crossveins; 8- CuA���A space expanded in middle to at least two cells wide, often more; 9- CuA long, terminating on posterior margin at mid-wing or longer (termination most proximal in Primorilestes species, but there mid-wing); 10- head width across eyes about twice the length from anterior margin of antefrons to posterior of occiput; compound eyes more or less adpressed to the head capsule, convex laterally but not spherical, posterolateral corners extended posteriorly to varying degrees, sometimes even acutely; distance between compound eyes at level of centre of ocelli about one eye���s width or less; head not distinctly extended laterally with bulging, spherical compound eyes as in Zygoptera (i.e., not ���hammerhead��� or ���dumbbell��� shaped). We agree with Nel & Arillo (2006), Nel & Fleck (2014), and Garrouste & Nel (2019) that an expanded CuA���A space is characteristic of the Dysagrionidae (character state 8). These character states associate Dysagrion, Phenacolestes, Primorilestes, Petrolestes, Electrophenacolestes, and Congqingia in agreement with previous authors, as well as Furagrion Petrulevičius et al. (restored, below); the new genera described here, Okanagrion, Okanopteryx and Stenodiafanus; and species of the new collective genus Dysagrioninites. They exclude Electrodysagrion, Palaeodysagrion and Burmadysagrion (see below). We tentatively include the Thanetian (late Paleocene: see age discussed by Wedmann et al. 2018) Valerea Garrouste et al. (one species: V. multicellulata, Menat Formation, France) for reasons discussed below. We include Furagrion in the Dysagrionidae and treat Valerea as a tentative member. Henriksen (1922) originally described Furagrion jutlandicus (Henriksen) from the early Ypresian Fur Formation of Denmark (���Mo-Clay���) as a species of Phenacolestes, and therefore a member of Cockerell���s Dysagrioninae. He based it on a specimen with three partial wings and part of the abdomen. None of these wings are preserved proximal to the nodus, and although they then lack the quadrangle and antenodal space, he associated the species with Phenacolestes by shared character states of the preserved portion. Owing to the lack of the proximal wing characters, Nel & Paicheler (1994) treated the species as ��� Phenacolestes ��� jutlandicus of indeterminate family. Rust (1999) reported a new, almost complete wing which he called ��� Dysagrioninae gen. indet. jutlandicus ���. Petrulevičius et al. (2008) revised the species based on this fossil, erecting the monotypic genus Furagrion for it. They assigned it to the Megapodagrionidae, finding an association with the Dysagrionidae unlikely by the lack of a distinct broadening of the antenodal area at the level of Ax1 and Ax2, and the distal side of the quadrangle not distinctly longer than the proximal. However, the amount of broadening of the antenodal area is variable in the family, and is in some species quite slight, and so this character state does not have clear diagnostic value in distinguishing it; in Furagrion it is comparable with that of Electrophenacolestes. While the distal side of the quadrangle of Furagrion is not much longer than the proximal side as it is in many dysagrionids, it is still longer, like that of Congqingia (Fig. 8). Furagrion then satisfies all wing character states of the diagnosis provided above, and so we restore it to the Dysagrionidae. Zessin (2011) described a second species of the genus, F. morsi Zessin, based on the proximal half of a wing. Although the enigmatic Thanetian Valerea multicellulata (Fig. 9) is known from a single, fragmentary wing in which only character state 1 of our family definition is determinable, we tentatively associate it with the Dysagrionidae by its otherwise notable similarity with the new genus Okanagrion with which it shares distinctive character states, some not otherwise known in the Odonata (see below, in the discussion of Okanagrion and support by our cladistic analysis). The Alaskan Chickaloon specimen and ��� Megapodagrionidae ��� genus and species A are probably dysagrionids; Thanetophilosina Nel et al. and NHMUK I.9866/I.9718 could be dysagrionids. We agree with Garrouste & Nel (2019) that the fragmentary specimen from the Thanetian or Ypresian Chickaloon Formation of Alaska (unnamed) is likely a dysagrionid by its density of veins and long CuA with an expanded CuA���A space up to four cells deep (Fig. 9). Its preserved portion only possesses character states 1, 8 and 9 of the Dysagrionidae diagnosis, and so we treat it here as a probable member of the family. Similarly, the partial wing ��� Megapodagrionidae ��� genus and species A of Petrulevičius et al. (2008) from the early Lutetian of Grube Messel, Germany, shares character states 1, 8 and 9 of the Dysagrionidae diagnosis with the Chickaloon specimen, and is more complete than it, with the apical portion present and well-preserved. Petrulevičius et al. (2008) discuss various possible family associations and only tentatively assign it to the Megapodagrionidae, noting that as then constituted, that family could not be defined by any known synapomorphies of wing venation. The Megapodagrionidae, long considered to be polyphyletic, has since been separated into a number of families based on a revised molecular phylogeny (Dijkstra et al. 2014). We treat this fossil as also probab, Published as part of Archibald, Bruce, Cannings, Robert A., Erickson, Robert J., Bybee, Seth M. & Mathewes, Rolf W., 2021, The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America, pp. 1-133 in Zootaxa 4934 (1) on pages 8-27, DOI: 10.11646/zootaxa.4934.1.1, http://zenodo.org/record/4558796, {"references":["Cockerell, T. D. A. (1908 a) Fossil insects from Florissant, Colorado. Bulletin of the American Museum of Natural History, 24, 59 - 69, 1 pl.","Calvert, P. P. (1913) The fossil Odonate Phenacolestes, with a discussion of the venation of the legion Podagrion Selys. Proceedings of the Academy of Natural Sciences of Philadelphia, 65: 225 - 272.","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. 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(2006) Dragonfly genera of the New World: an illustrated and annotated key to the Anisoptera. Johns Hopkins University Press, Baltimore, Maryland, 368 pp.","Tillyard, R. J. & Fraser, F. C. (1940) 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 III, suborder Anisozygoptera. Australian Zoologist, 9, 359 - 396.","Hagen, H. (1858) Zwei Libellen aus der Braunkohle von Sieblos. Paleontographica, 5, 121 - 124.","Huang, D., Fu, Y. & Nel, A. (2019) The first amber stenophlebiid damsel-dragonfly (Odonata, Epiproctophora, Stenophlebiidae) from mid-Cretaceous amber of northern Myanmar. Cretaceous Research, 94, 40 - 44. https: // doi. org / 10.1016 / j. cretres. 2018.10.008"]}

Okanagrion beardi Archibald & Cannings, new species Figs. 21–22. Diagnosis. Wings separated from those of all other Okanagrion species but O. dorrellae by MP, CuA distinctly divergent near margin [others: MP-CuA space remaining narrow; O. dorrellae: widening, but less distinctly as CuA zigzagged distally (O. angustum, not known)]; from all others except O. angustum by IR2 originating on RP1-2 [others: between RP1-2, RP3-4]; from O. angustum by Ax1 to nodus shorter, 6.5 mm (holotype; paratype 1: almost 6 mm, smaller wing, but proportions match holotype) [O. angustum: 7.5 mm]; from O. dorrellae by MP linear to margin [O. dorrellae: zigzagged near margin]; from O. liquetoalatum, O. dorrellae by wing shape: length (arculus to apex) / maximum width 3.1 [O. liquetoalatum: 2.6; O. dorrellae 3.7; others similar to O. beardi (O. lochmum, O. worleyae, O. angustum not comparable by preservation)]. Further separated by colouration (sex unknown): proximal approx. 2/5 hyaline, dark apical to this except narrow lighter fascia just distal to mid-way between nodus, pterostigma, extending from anterior margin to close to posterior margin; size and extent of light fascia differs from: O. hobani about twice or more as wide, extending to or almost to posterior margin; O. angustum apparently similar to O. hobani as preserved; O. threadgillae notably narrower, extending to mid-wing; O. dorrellae similar width, probably extending to mid-wing (but damage in this region); O. lochmum, similar to O. beardi; O. liquetoalatum: hyaline throughout; O. worleyae: lightly infuscate throughout known wing. Type material. Holotype: F-791 (part only) (Fig. 21): One complete wing and portions of two or three others; possibly part of one eye; part of damaged thorax preserved in lateral aspect; at least two pieces of abdomen present; collected at the McAbee Hoodoo Face beds, collector and date unknown; housed in the Thompson Rivers University collections. Paratype 1: RBCM P1546 (Fig. 22): a head, thorax, bits of some legs, proximal-most portion of the abdomen, well-preserved forewing and proximal portion of a hind wing; coated with shellac; collected at the McAbee Hoodoo Face beds, vi.1998 by unknown grade 8 or 9 student, donated by Graham Beard; in the Royal British Columbia Museum collections. Description. Holotype, F-791, forewing. Almost complete, missing tiny portions of the apex and proximal to the nodus. Shape: oval distal to nodus, symmetrical from lateral mid-line; length: 31.5 mm, arculus to apex: about 27.7 mm, nodus to apex: about 22.7 mm, origin of RP2 to apex: 18.2 mm, arculus to base of pterostigma: 22.0 mm, nodus to base of pterostigma: 16.8 mm, nodus at 29% wing length; width 8.3 mm. Colouration as in diagnosis. Pterostigma anterior, posterior margins oblique, 3.5 times longer than wide, subtends 6.5 cells, no oblique brace vein. 35 crossveins in postnodal space (complete), none hyperstigmal, crossveins in postsubnodal space not aligned. Maximum three cells wide distal to pterostigma in costal space (partially preserved). RA–RP1 space narrows near margin. RP1–IR1 space becomes two cells wide unknown number of cells from origin by preservation. IR1 origin eight cells distal to origin of RP2. IR1–RP2 space becomes two cells wide unknown number of cells origin by preservation. RP2 origin six cells distal to origin of IR2. IR2 origin very close to subnodus; close to, appears to arise from RP1-2. RP3-4 origin about 2/3 from arculus to nodus. Ax0 present. Arculus close to, immediately proximal to Ax2. No crossvein O. All main veins linear other than IR1 proximally. MP, CuA divergent at terminus on posterior margin. CuA–A space with maximum five cells wide. Paratype 1, RBCM P1546. Head (Fig. 10 M, Table 1): relatively well preserved in dorsal aspect but details obscure; labrum/clypeus area damaged. In dorsal view about 4.1 mm across eyes at widest part, with rear margin expanded somewhat postero-laterally; approximate length from anterior edge of antefrons to posterior of occiput 1.5 mm; at level of hind ocelli distance between eyes about 1.2 mm, single eye roughly 1.5 mm wide, 2.0 mm long. Thorax: lateral view; colouration dark, apparently with narrow pale stripe on mesopleural suture, any other patterns obscure; pieces of legs poorly preserved. Forewing: as in holotype except length: 31.6 mm, arculus to apex: 26.5 mm, nodus to apex: 21.7 mm, origin of RP2 to apex: 16.6 mm, arculus to base of pterostigma: 21.0 mm, nodus to base of pterostigma: 16.2 mm; width 7.7 mm; nodus at 25% wing length. Colouration mostly obscured by applied shellac, but as apparent is consistent with holotype. Pterostigma: 5.5 cells subtend, length 3.5 times width. Postnodal space with 36 crossveins (probably complete), distal-most one hyperstigmal. IR1 origin seven cells distal to origin of RP2; IR1– RP2 space becomes two cells wide six or seven cells (unclear by preservation) distal to origin by preservation. RP2 origin nine cells distal to origin of IR2. Ax0 not detected. CuA–A space unknown number of cells wide by preservation, but appears similarly wide as holotype. Abdomen: segments 1 and 2 preserved in lateral aspect; base of abdomen relatively slender; irregular ventral margin of segment 2 may indicate presence of secondary male genitalia but sex of specimen undetermined. Etymology. The specific epithet is a patronymic formed from the surname of Graham Beard, the donor of paratype 1, recognising his contribution. Range and age. McAbee, BC, Canada; mid-Ypresian.

Okanopteryx incertae sedis Figs. 58–60. Material. Republic: SRUI 99-72-96AB (Fig. 58), collected at Republic B 4131 by Rita Weikel, 27.iv.2019, housed in the Stonerose Interpretive Center collections; SRUI 99-97-44 (Fig. 58), collected at Republic A0307 by Wes Wehr, date unknown, in the Stonerose Interpretive Center collections; F-1046 (Fig. 58), collected at McAbee, collector and date unknown, in the Thompson Rivers University collections; UWBM-72324 (Fig. 60), collected by Wes Wehr at Republic A0307, date unknown, in the Burke Museum collections; SR 13-005-015 AB (Fig. 59), collected at Republic B 4131 by Richard Dillhoff, 6.v.2008, in the Stonerose Interpretive Center collections; GSC 141103 (Fig. 59), collected at the McAbee Hoodoo Face beds, Geological Survey of Canada locality code V-016800, by SBA, July 31 2000, in the Geological Survey of Canada collections; RBCM P1553 (Fig. 60), collected at the McAbee Hoodoo Face beds by John Leahy, date unknown, in the Royal British Columbia Museum collections. Discussion. These wing fragments of varying size appear to belong to one of the defined species of Okanopteryx by their distinctive colouration and all other preserved morphology, but diagnostic character states separating these species are not preserved. For example, F-1046 agrees with the O. macabeensis holotype in all aspects, e.g., just over 14 mm nodus to pterostigma, but lacks almost the entire CuA–A space. The preserved region of specimen SR 13-005-015 AB is almost identical to that of the more complete O. jeppesenorum SR 02-32-23, also from Republic exposure B4131.

Okanagrion worleyae Archibald & Cannings, new species Figs. 39–41. Diagnosis. Wings separated from those of O. threadgillae, O. beardi, O. liquetoalatum, O. lochmum, O. hobani, by shape: posterior margin less deeply curved; further from O. threadgillae by shape, nodus to apex 3.0 times maximum width [O. threadgillae: 2.5]; further from O. angustum by narrower wing, about 4.5 mm wide at levels of nodus [O. angustum about 5.5 mm]; from O. dorrellae by MP linear near margin (but partially known by preservation) [O. dorrellae: zigzagged]; from O. beardi, O. dorrellae by MP, CuA sub-parallel at terminus on posterior margin [O. beardi: distinctly wider, O. dorrellae: widening, but less distinctly as CuA zigzagged distally (O. angustum, not known); from O. angustum, O. beardi by IR2 origin between RP1-2, RP3-4 (clearly preserved in holotype, not clear by damage in paratype 2, region not preserved in paratype 1) [O. angustum, O. beardi: origin on RP1-2]; from O. hobani by main veins less deeply curved near margin; from O. lochmum by cells less dense in apical portion of wing: RA–RP1 space two cells wide distal to four cells proximal to pterostigma [O. lochmum: 7–8 cells proximal], RA–RP1 space three cells wide about one cell distal to pterostigma [O. lochmum: at base of pterostigma], RA–RP1 space maximum four wide from about four cells distal to pterostigma [O. lochmum: four, then five cells wide about distal corner of pterostigma]. Further separated by colouration (sex unknown): wing lightly, uniformly infuscate across all known portions as preserved (might appear almost hyaline by preservation). Type material. Holotype: SR 99-14-02 (Fig. 39), collected at Republic A0307 by Providence Worley, 23.viii.1998, housed in the Stonerose Interpretive Center collections; paratype 1: SR 16-006-001 (Fig. 40), collected at Republic A0307 by Kattia Rojas, 9.x.2016, in the Stonerose Interpretive Center collections; paratype 2: SR 11-21-09 (Fig. 41), collected at Republic B 4131 by Eric Blumhagen, 2.viii.2008, in the Stonerose Interpretive Center collections. Description. Holotype, SR 99-14-02, wing (Fig. 39). Missing portion proximal to nodus. Shape: narrow oval distal to nodus, mostly symmetrical from lateral mid-line, except apex slightly anterior to this. Length: nodus to apex: 22.7 mm, origin of RP2 to apex: 18.5 mm, nodus to base of pterostigma: 17.2 mm; width 7.8 mm. Lightly infuscate throughout. Pterostigma anterior, posterior margins oblique, 3.5 longer than wide, subtends seven cells, no oblique brace vein. 37 crossveins preserved in postnodal space (probably two not preserved), distal-most one hyperstigmal; those in postsubnodal spaces not aligned. Three cells wide distal to pterostigma in costal space. RA meets margin close to, posterior to apex, RA slightly curved forward near margin. RA–RP1 space narrows at margin. RP1–IR1 space goes to two cells wide nine cells distal to origin of IR1. IR1 origin eight cells distal to origin of RP2; IR1 two cells wide eight cells distal to origin. RP2: origin apparently (by preservation) seven cells distal to origin of IR2. IR2 origin at subnodus. No crossvein O. All major veins linear except proximal portion of IR1. CuA–A space four cells wide at widest. Paratype 1, SR 17-006-001, wing (Fig. 40). Apical portion, from about termination of MA on posterior margin to apex; diagonally to near nodus on anterior margin. As for holotype except: pterostigma subtends 4.5 cells; four cells wide distal to pterostigma in costal space; RA terminates a bit further posteriad. Paratype 2, SR 11-21-09, wing (Fig. 41). Preserved from about level of nodus, missing portion of posterior-apical region, partly indistinctly preserved.As in holotype, except: length: nodus to apex: about 22.7 mm, origin of RP2 to apex: 17.9 mm, nodus to base of pterostigma: about 17.2 mm. Pterostigma almost four longer than wide, subtends unknown number of cells. Crossveins in postnodal space partially preserved, appear as dense as in holotype, none hyperstigmal. RA not preserved at margin. RP1-2 –IR2, RP1 –IR1, IR1– RP2, IR2– RP3-4 spaces partially preserved, IR1, RP2 origin distances appear similar to holotype. IR2 origin at subnodus. No crossvein O. All major veins linear except proximal portion of IR1. CuA–A space five cells wide at widest (but region poorly preserved). Etymology. The specific epithet is a patronymic formed from the surname of Providence Worley, the collector and donor of the holotype, recognising her contribution. Range and age. Exposures B 4131 and A0307 of the Tom Thumb Tuff Member of the Klondike Mountain Formation at Republic, Washington, USA; late Ypresian., Published as part of Archibald, Bruce, Cannings, Robert A., Erickson, Robert J., Bybee, Seth M. & Mathewes, Rolf W., 2021, The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America, pp. 1-133 in Zootaxa 4934 (1) on pages 70-75, DOI: 10.11646/zootaxa.4934.1.1, http://zenodo.org/record/4558796

Stenodiafanus westersidei Archibald & Cannings, new species Fig. 61. Diagnosis. As for genus. Material. Holotype: SR 06-01-42, collected at Republic B 4131 by Steven Westerside, 15.viii.2005, housed in the Stonerose Interpretive Center collections. Description. Holotype, sex unknown. SR-06-01-42, wing (Fig. 61). Hyaline, narrow; nodus to apex 24.8 mm, width 5.8 mm. 18 postnodal crossveins, about half aligned with postsubnodal crossveins, mostly the proximal third. Pterostigma pale, four times longer than wide. Costal space apical to pterostigma one cell wide for one cell, then two. Oblique brace vein present, connecting to proximal corner of pterostigma, oblique. RA–RP1 space one cell wide except for a few adventitious crossveins. RP1–IR1, RP2–IR2, IR2–RP3-4 spaces become two cells wide about half distance from origin of IR1 to pterostigma; IR1–RP2 space becomes two cells wide shortly distal to this. RA, RP1, IR1, RP2 converge, almost meeting at apex (only six cells likely along margin between RA, RP2). RP2 origin about quarter distance between nodus, pterostigma. IR2: origin at level of subnodus, slightly zigzagged in distal quarter. RP3-4: origin proximal to, near nodus; rather straight from level of nodus to terminus on margin at level of distal end of pterostigma. MA linear from level of nodus (not preserved proximal to that) to about half way to level of pterostigma, then zigzagged to margin, about 3-4 nodus to pterostigma levels. MP preserved from level of nodus: rather straight to slight curve near terminus on margin. CuA preserved from just proximal to origin of RP2, zigzagged to margin. Etymology. The specific epithet is a patronymic formed from the surname of Steven Westerside, the collector and donor of the holotype, recognising his contribution. Range and age. Tom Thumb Tuff Member of the Klondike Mountain Formation, locality B4131 at Republic, Washington, USA; late Ypresian.

Whetwhetaksidae Archibald & Cannings, new family Figs. 66–70. Diagnosis. Most easily separated from all other Cephalozygoptera and Zygoptera families by 1, distinctively long pterostigma: length about ten times width (roughly, where known), nodus to base of pterostigma 55–60% of nodus to apex; and from all Cephalozygoptera and all Zygoptera except the extant Heliocharis amazona (Dicteriadidae) and some Paleogene Euphaeidae by 2, arculus just distal to Ax1, closer to it than to Ax2. Differs further from Sieblosiidae by: 3, oblique vein O absent [present]; 4, nodal, subnodal crossveins oblique [usually nearly vertical, sometimes slightly oblique or reversed obliquity]; 5, base of RP2 close to nodus [more distal]; 6, CuA terminates at margin over half wing length [notably shorter, longest Parastenolestes oligocenicus Nel & Paicheler, under half wing length]; 7, MA close to, subparallel with RP3-4 to margin [subparallel with MP]; 8, MA, MP diverge to margin [remain close]; 9, MP, CuA remain subparallel to margin [widely diverge]; 10, base to nodus about 40% wing length, minimum about 38% [maximum about 35% where measurable]. Differs further from Dysagrionidae by: 4 [usually like Whetwhetaksidae, but Dysagrion subnodal crossvein reverse obliquity]; 7–9, [varies], 10, [Dysagrion packardii, about 42%, Petrolestes hendersoni, about 42%, the rest about 36–25%]; 11, origin of IR 2 in middle third between arculus, nodus [at subnodus in Dysagrioninae, but as in Whetwhetaksidae, in Petrolestinae]. Type and included genera. Whetwhetaksa. Description. As for its only genus, see below. Etymology. The family name is formed from that of its type and only genus, Whetwhetaksa. Range and age. Republic localities B4131 and A0307B of the Tom Thumb Tuff Member of the Klondike Mountain Formation; latest Ypresian. Discussion. Such an extremely long pterostigma is never found in the Zygoptera to our knowledge, but is matched by some Petaluridae (Anisoptera). The position of the arculus closer to, immediately distal to Ax1 is remarkable. This condition is found in the Anisozygoptera and Anisoptera, but in the Zygoptera only in the South American H. amazona, which otherwise differs strongly in many ways and a few Paleogene Euphaeidae (see above) which also strongly differ. In all other Zygoptera it is aligned with, slightly proximal to, or slightly to clearly distal to Ax2 (Bechly 1996; Fleck et al. 2004). In SR 06-01-36, the subcosta appears to cross through the nodus, which is characteristic of the Sieblosiidae (see, e.g., Nel & Fleck 2012, their Fig. 3). This is not present in any other specimen of Whetwhetaksa, and it could be a crossvein adventitiously positioned between the costa and nodus in this specimen and not the subcosta. We tentatively associate the Whetwhetaksidae with the Cephalozygoptera by the many similarities of their wings (see description of the suborder, above). The wings of Whetwhetaksidae are more like those of the Cephalozygoptera than any Zygoptera, but await confirmation of this relationship by the discovery of a more complete fossil that includes the head., Published as part of Archibald, Bruce, Cannings, Robert A., Erickson, Robert J., Bybee, Seth M. & Mathewes, Rolf W., 2021, The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America, pp. 1-133 in Zootaxa 4934 (1) on page 110, DOI: 10.11646/zootaxa.4934.1.1, http://zenodo.org/record/4558796, {"references":["Bechly, G. (1996) Morphologische Untersuchungen am Flugelgeader der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata) unter besonderer Beruchsichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata. Petalura, Boblingen, Special Volume, 2, 1 - 402.","Nel, A. & Fleck, G. (2012) A new genus and species of Sieblosiidae from the middle Miocene of Germany (Odonata: Epiproctophora). Zootaxa, 3582 (1), 64 - 68. https: // doi. org / 10.11646 / zootaxa. 3582.1.7"]}

Dysagrionites delinei Archibald & Cannings, new species Figs. 62–63A. Diagnosis. Wings of Dysagrionites delinei may be distinguished from those of all other species of Dysagrioninae except of Okanopteryx, Stenodiafanus by: Cu–A space 3 cells wide [Okanagrion: 4–7; Furagrion, Dysagrionites sp. A, Stenodiafanus: 2 (most likely, wing incomplete); Dysagrion, Phenacolestes: 4; Electrophenacolestes, 4 (one column 5); Okanopteryx: 2–4; not separated from Primorilestes: 3–5; Dysagrionites sp. B: probably 3 but unclear]. By CuA zigzagged beyond about level of origin of RP2 [Electrophenacolestes, Primorilestes, Okanagrion: linear; not separated from Dysagrion, Okanopteryx, Phenacolestes, Stenodiafanus]. Separated from the sole species of Stenodiafanus by wider wing, 6.5 mm wide [5.8 mm]; by CuA–A space maximum three cells wide [two]. Separated from males of Okanopteryx, Okanagrion (males of O. hobani, sex unknown in other species) except O. liquetoalatum, Dysagrionites species B (sex unknown) by wing hyaline throughout known portion (from base past presumed mid-way between nodus, pterostigma [Okanopteryx, Okanagrion except O. liquetoalatum, Dysagrionites species B: with infuscate regions]. Further distinct from Primorilestes by no intercalary vein in CuA–A space. Most easily separated from fossil genera regarded here as possible Dysagrionidae by: from Chickaloon specimen by cells of CuA–A evenly sized, spaced, maximum three wide [irregularly sized, spaced, four wide]; Thanetophilosina by all preserved crossveins in postnodal, postsubnodal spaces aligned or almost so [not]; from NHMUK I.9866/I.9718 by origin of RP3-4 about a third distance from subnodus to level of anterio-distal corner of quadrangle [about 20%]; distinct from cf. Dysagrionidae genus A, species A (Petrulevičius et al. 2008) by hyaline membrane [infuscate except apex distal to pterostigma]; CuA lightly zigzagged distal to level of origin of RP2 [linear]. Material. Holotype: SR 06-01-46AB (Fig. 62), collected at Republic B 4131 by William Deline, 21.ix.2005, housed in the Stonerose Interpretive Center collections; paratype 1: UWBM 77612 (Fig. 63A), collected at Republic, no exposure, collector, or date recorded, in the Burke Museum collections; paratype 2: SR 94-05-26 (Fig. 62), collected at Republic B 4131 by Analee Flowers, 18.x.1994, in the Stonerose Interpretive Center collections. Description. Holotype, SR 06-01-46, wing. Missing apical portion from proximal to pterostigma. Approximately 6.5 mm maximum width. Hyaline throughout (sex unknown). All preserved crossveins in postnodal, postsubnodal spaces aligned or almost so. RP1–IR1, IR1–RP2 spaces do not become two cells wide in preserved portion. IR1: origin four cells distal to origin of RP2. RP2: origin almost five cells distal to origin of IR1. IR2: origin at subnodus. RP3-4: origin not preserved. MA with only slight zigzag around level of origin of IR1. MP linear, slightly curved from quadrangle (distal portion not preserved). CuA becomes slightly zigzagged around level of nodus, increasingly so apical to level of origin of RP2. CuA–A space three cells at widest (from close distal to level of nodus to level of origins of IR1), 9–10 columns; two cells wide proximal to this about 4 columns, distal to this a few columns of two cells wide, then not preserved. Paratype 1, UWBM-77612, wing. Proximal portion missing apical portion from mid-wing. Maximum width not measurable, but preserved portions match SR 06-01-46. Hyaline throughout (sex unknown). Three preserved crossveins in proximal postnodal, postsubnodal spaces aligned. RP1 –IR1, IR1– RP2 spaces do not become two cells wide in preserved portion. IR1 not preserved. RP2: origin probably five cells distal to origin of IR1. IR2: origin at subnodus. RP3-4: origin just over 2/3 distance from arculus to subnodus. MA with only slight zigzag around similar length as SR 06-01-46. MP linear, slightly curved from quadrangle (distal portion not preserved). CuA becomes slightly zigzagged around level of nodus, increasingly so apical to approximate level of origin of RP2. CuA–A space two cells wide from level of nodus to half way between level of nodus, presumed origin of RP2; three cells wide 6–7 columns, then briefly two for remaining preserved portion. Paratype 2, SR 94-05-26, wing. Anterior portion missing apical region proximal region, much of posterior region. Hyaline throughout (sex unknown). All preserved crossveins in postnodal, postsubnodal spaces aligned or almost so, two in nodal space without matching subnodal space crossveins. RP1 –IR1 space becomes two cells wide six cells distal to origin of IR1. IR1: origin three cells distal to origin of RP2; IR1– RP2 space does not become two cells wide in preserved portion. RP2: origin apparently 3.5 cells distal to origin of IR1. IR2: origin at subnodus. RP3-4: origin 2/3 length of arculus to subnodus. MA notably zigzagged beyond level of origin of IR1. MP: small portions preserved, linear. CuA slightly zigzagged around level of origin of RP2, then increasingly so in remaining preserved portion. CuA–A space at least three cells wide at least seven columns, maybe (many?) more, all preserved portion at least two. Etymology. The specific epithet formed from the surname of William Deline, collector and donor of the holotype, recognising his contribution. Range and age. Tom Thumb Tuff Member of the Klondike Mountain Formation at Republic, Washington, USA, all with exposure information are from B4131; late Ypresian. Discussion. This species might belong to either Okanopteryx or Okanagrion, expanding its genus concept, or may constitute a new orthotaxon genus. These differ among themselves by the width of the CuA–A space: SR 06-01-46 is three cells wide between the origins of RP2 and IR1, SR 96-12-03 is three cells wide only briefly around the level of the origin of RP2, and in UWBM-77612 it is uniformly two cells wide except for one column of three. This suggests that there may be more than one species among these specimens. Although SR 96-12-03 is the middle portion of a wing lacking almost all diagnostic character states of the Dysagrionidae listed above, its great similarity to the other two specimens in particular makes it possible to assign it to the family. It does differ in having almost the whole preserved CuA–A space two cells wide, whereas in the other specimens it has many columns of three. If this specimen is not a member of this species, we believe it likely closely related.