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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"]}

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 (���������), 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 568, DOI: 10.11646/zootaxa.5100.4.6, http://zenodo.org/record/6225002

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 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

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 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.

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.

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., 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 102, DOI: 10.11646/zootaxa.4934.1.1, http://zenodo.org/record/4558796

Dysagrionites sp. A Fig. 63B. Material. SR 96-12-03 (Fig. 63B), collected at Republic B 4131 by Les Larson, 15.x.1996, housed in the Stonerose Interpretive Center collections. Description of SR 96-12-03. The mid-portion of wing apical to about origin of RP2 to shortly distal to origin of IR1 anteriorly diagonally to shortly proximal to terminus of RP3-4. Width estimated 6.5 mm. hyaline throughout. Few preserved crossveins in proximal postnodal, postsubnodal spaces aligned. RP1–IR1 space does not become two cells wide in preserved portion. IR1: origin three cells distal to that of RP2; RP2–IR2 space becomes two cells wide ten cells distal to origin. RP2 linear, rather straight through preserved portion. IR2, RP3-4 origins not preserved, rather linear, slightly zigzagged in places, rather straight through preserved portion. MA increasingly zigzagged through preserved portion. MP linear, slightly curved in portion preserved to near terminus. CuA zigzagged throughout preserved portion. CuA–A space mostly preserved, from just proximal to level of origin of RP2 almost to distal end; two cells wide throughout with one column of three. Range and age. Okanagan Highlands, Thumb Tuff Member of the Klondike Mountain Formation at Republic, Washington, USA, exposure B4131; late Ypresian. Discussion. This middle portion of a wing most resembles Dysagrionites delinei in that it is narrow, hyaline, three cells from origin of RP3-4 to origin of IR1 [D. delinei 3–4], CuA linear to mid-wing, then begins to zigzag (distal portion not known). It differs, however in the CuA–A space being two cells wide (one column three), whereas in D. delinei it is three cells wide. It differs from Furagrion and hyaline species of Okanagrion by CuA zigzagged in distal portion to the margin, whereas in Furagrion it is lightly zigzagged in mid-wing, returning to linear in the distal portion to the margin, and in Okanagrion it is linear throughout. It differs further from hyaline Okanagrion species by the narrow CuA space [four or more cells wide]. Although it is distinct in these ways, we prefer to wait to name this species until a more complete specimen is found that would allow a more through definition, and given the chance of sexual dimporphism in wing colouration., 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 106, DOI: 10.11646/zootaxa.4934.1.1, http://zenodo.org/record/4558796

Okanopteryx jeppesenorum Archibald & Cannings, new species Figs. 44–48. Diagnosis. Wings distinct from those of O. macabeensis, O. fraseri by CuA–A space wider, four cells wide at widest (difficult to count number of columns by preservation), between the levels of origins of IR1, RP2 [O. macabeensis: 5–10 columns of three cells wide; O. fraseri: all two cells wide, but sometimes a single column of three]; further from O. macabeensis, by longer wing, holotype, 22.4 mm nodus to apex; paratype 1 (SR 11-43-09): 23.5 mm [O. macabeensis holotype: estimated 18.4 mm] (all O. fraseri wings missing apex). Material. Holotype: SR 13-005-013 (Fig. 44), collected at B4131 by Jackie and Tim Jeppesen, 15.vii.2013, housed in the Stonerose Interpretive Center collections; paratype 1: SR 11-43-09, (Fig. 45), collected at Republic B 4131 by Christin Gustafson, 15.vi.2011, in the Stonerose Interpretive Center collections; paratype 2: SR 18-002- 001 (Fig. 46), collected at Republic B 4131 by Valerie Paquin, 28.iv.2018, in the Stonerose Interpretive Center collections; paratype 3: SR 15-003-001 (Fig. 47), collected at Republic B 4131 by Ethan Fisher, 30.v.2015, in the Stonerose Interpretive Center collections; paratype 4: SR 02-32-23 (Fig. 48), collected at Republic B 4131 by Chris Engle, 6.vi.2002, in the Stonerose Interpretive Center collections; paratype 5: SR 07-22-05AB (Fig. 47), collected at Republic B 4131 by Kathleen Malloch 13.viii.2006, in the Stonerose Interpretive Center collections; paratype 6: SR 19-005-002AB (Fig. 48), collected at Republic B 4131 by Gregg Wilson, 1.v.2019, in the Stonerose Interpretive Center collections. Description. Holotype, SR 13-005-013, wing (Fig. 44). Mostly complete, with portions of mid-posterior not preserved. Petiolate. Evenly oval distal to nodus, nodus to apex 22.4 mm, maximum width mid-wing 7.7 mm. Hyaline except for broad dark fascia across wing between level of nodus, just over half way to pterostigma. Pterostigma broad, about 2.5 times longer than wide, proximal side nearly vertical; subtends 2.5 cells. Oblique brace vein joins RA at proximal corner of pterostigma. Most crossveins roughly aligned in proximal half of postnodal, postsubnodal spaces. Costal space distal to pterostigma two cells wide. RA meets margin posterior to apex, RA slightly sigmoidal near margin. RA–RP1, RP1–IR1 spaces narrow near margin. RA–RP1 space mostly one cell wide, briefly two near apex. RP1–IR1 space two cells wide four cells distal to origin of IR1. Origin of IR1 presumed four cells distal to origin of RP2, mid-way between arculus, pterostigma. IR1–RP2, RP2–IR2 spaces two cells wide shortly proximal to level of pterostigma. Origin of RP2 about 5–6 cells distal to subnodus. RP2 mostly rather straight, curved slightly posteriorly in distal quarter. IR2: origin at subnodus; mostly rather straight, linear, slightly zigzagged in distal quarter, strongly so near margin. Origin of RP3-4 about 2/3 between arculus, nodus; mostly rather straight. MA linear, straight at least to half way from quadrangle (partially not preserved), strongly zigzagged at least distal third. MP linear, rather straight from quadrangle to terminus. CuA linear until shortly after level of nodus, then increasingly zigzagged. CuA–A space with 4 cells at widest (from just proximal to, distal to level of RP2 origin as preserved). Paratype 1, SR 11-43-09, wing (Fig. 45). Petiolate. Evenly oval distal to nodus; nodus to apex 23.5 mm: maximum 7.2 mm wide. Hyaline except for broad dark fascia across wing between level of nodus, just over half way to pterostigma. Pterostigma broad, about 2.5 times longer than wide; proximal side with shallow angle to costa. Region of oblique brace vein not preserved. Crossveins in postnodal, postsubnodal spaces mostly rather aligned. Two cells wide distal to pterostigma in costal space. RA–RP1 space one cell wide. RA meets margin posterior to apex. RP1 –IR1 space goes to two cells wide five cells distal to origin of IR1; narrows near margin. Origin of IR1 four cells distal to origin of RP2, mid-way between arculus, pterostigma. IR1– RP2, RP2 –IR2 spaces mostly not preserved. RP2: origin about 3 cells distal to subnodus; proximal half linear, rather straight; distal half not preserved. IR2: origin at subnodus; mostly rather straight, liner, slightly zigzagged in distal quarter, missing by preservation near margin. RP3-4: origin Paratype 2, SR 18-002-001, wing (Fig. 46). Petiolate. Evenly oval distal to nodus (missing apical region), nodus to base of pterostigma 16.5 mm, maximum width mid-wing 7.2 mm. Hyaline except for broad dark fascia across wing between level of nodus, to about presumed 2/3 distance to pterostigma. Pterostigma not preserved. Most crossveins roughly aligned in proximal half of postnodal, postsubnodal spaces, not in distal half. RP1 –IR1 space two cells wide eight cells distal to origin of IR1. IR1 origin two cells distal to origin of RP2. RP2: origin apparently five cells distal to subnodus; mostly rather straight in region preserved. IR2: origin at subnodus; mostly rather straight, linear, apical quarter missing. RP3-4: origin missing; mostly rather straight, missing apical quarter. MA linear, straight at least to half way between nodus, presumed level of pterostigma. MP linear, rather straight from quadrangle to terminus. CuA linear until shortly after level of nodus, then slightly zigzagged to distal quarter, then increasingly so. CuA–A space four cells at widest from just proximal to level of origin of RP2 to about level IP2 origin. Paratype 3, SR 15-003-001, wing (Fig. 47). Proximal two-thirds. Petiolate. Maximum width 7.7 mm. Hyaline proximal to level of nodus, darkly infuscate distal to it. Most crossveins roughly aligned in proximal half preserved of postnodal, postsubnodal spaces. RP1 – IR1 space two cells wide five cells distal to origin of IR1. IR1: origin four cells distal to origin of RP2. RP2: origin five cells distal to subnodus; mostly rather straight in region preserved. IR2: origin at subnodus; mostly rather straight, linear, apical half missing. RP3-4: origin missing; mostly rather straight, missing apical third. MA mostly linear, slightly zigzagged around levels of origin of RP2 to IR1, mostly not preserved distal to that. MP mostly linear, slightly zigzagged in portions, gently curved from quadrangle, near terminus not preserved. CuA rather linear to level of origin of RP2, then increasingly zigzagged. CuA–A space four cells at widest from just proximal to, to just distal to level of origin of RP2. Paratype 4, SR 02-32-23, wing (Fig. 48). Apical half, missing much of anterior region. Apical portion hyaline from about half way between origin of IR1, pterostigma; darkly infuscate proximal to this. Pterostigma broad, proximal portion not preserved. Postnodal, most of postsubnodal spaces not preserved. Costal space distal to pterostigma two cells wide. RA meets margin posterior to apex. RA–RP1, RP1 –IR1 spaces narrow near margin. RA–RP1 space mostly one cell wide, briefly two near apex. RP1 –IR1 space constantly two cells wide twelve cells distal to origin of IR1. Origin of IR1 presumed three cells distal to origin of RP2. IR1– RP2 space two cells wide at presumed level of proximal side of pterostigma; IR2– RP3-4 space two cells wide from a few cells proximal to this. RP2: linear, mostly rather straight, slightly curved posteriorly in apical quarter. IR2 straight, linear to level RP2 –IR2 space becomes two cells wide, then slightly zigzagged, curves posteriorly near terminus. RP3-4: mostly rather straight, wing damaged near margin. MA preserved portion linear to about level of origin of IR1, then increasingly zigzagged. MP preserved portion linear, slightly curved to terminus. CuA increasingly zigzagged in region preserved, strongly so apically. CuA–A space four cells wide from just proximal to level of RP2 origin to just distal to level of origin of IR1. Paratype 5, SR 07-22-05 A, B, wing (Fig. 47). Parts of proximal half. Petiolate. Hyaline proximal to level of nodus, darkly infuscate distal to it. Most crossveins in postnodal, postsubnodal spaces roughly aligned in region preserved. Origin of IR1 three cells distal to origin of RP2. RP2: origin 2.5 cells distal to subnodus; portion preserved rather straight, linear. IR2: origin at subnodus; straight, linear. RP3-4, MA, MP: origins, distal portions missing; preserved portion mostly rather straight, rather linear. CuA–A space 4 cells wide just proximal to level of RP2 origin, not preserved distal to this. Paratype 6, SR 19-005-002 A, B, wing (Fig. 48). Broken up middle portion from closely distal to nodus to base of pterostigma, posterior region missing or damaged. Proximal portion darkly infuscate, distal portion hyaline. Many crossveins roughly aligned in preserved portion of postnodal, postsubnodal spaces. IR1 origin four cells distal to origin of RP2. RP2, IR2, RP3-4, MA: preserved portions mostly linear, rather straight, except MA zigzagging in distal portion preserved. Small portions of MP reserved. Disarticulated fragment of CuA–A space preserved, four cells wide. Etymology. The specific epithet is a patronymic formed from the surname of Jackie and Tim Jeppesen, the collectors and donors of the holotype, recognising their contribution. The Latin suffix – orum indicates the genitive case of plural nouns. Range and age. Tom Thumb Tuff Member of the Klondike Mountain Formation, locality B4131 [and other maybe?] at Republic, Washington, USA; second half of the Ypresian.