Okanopteryx macabeensis Archibald & Cannings, 2021

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, Zootaxa 4934 (1), pp. 1-133 : 91-97

Okanopteryx macabeensis Archibald & Cannings , new species

Figs. 53–57 .

Diagnosis. Wings distinct from those of O. fraseri , O. jeppesenorum as in their diagnoses; further from O. fraseri by abdomen/body length as in its diagnosis.

Material. Holotype: GSC 141100 (collector number SBA-2801) ( Fig. 53 ), collected at the McAbee Hoodoo Face beds, Geological Survey of Canada locality code V-016800, by SBA, 10.vii.2000, housed in the Geological Survey of Canada collections; paratype 1: GSC 141101 ( Fig. 54 , 55A, 55 C–55F), collected at the McAbee Hoodoo Face beds, Geological Survey of Canada locality code V-016801 by Rod Bartlett, vii.1991, in the Geological Survey of Canada collections; paratype 2: F-790 ( Fig. 56 ), collected at the McAbee Hoodoo Face beds, date and collector unknown (this is the much less complete counterpart; the part, with the almost entire insect preserved, was last seen in the inaccessible private collection of a commercial fossil dealer), in the Thompson Rivers University collections; paratype 3: GSC 141102 (collector number SBA-5928) ( Fig. 57 ), collected at the McAbee Hoodoo Face beds, Geological Survey of Canada locality code V-016800, by SBA, date unknown, in the Geological Survey of Canada collections; paratype 4: RBCM 3087 ( Fig. 53 ), collected at the McAbee Hoodoo Face beds by John Leahy , date unknown in the Royal British Columbia Museum collections.

Description. Holotype, GSC 141100, wing ( Fig. 53 ). Almost complete, missing petiole, small portion of the posterior margin in the apical quarter. Oval distal to nodus. Nodus to apex estimated 18.4 mm (presumed: small portion of apex missing), nodus to pterostigma 14.2 mm; width 6.5 mm wide. Hyaline except for broad, darkly infuscate fascia from level of nodus to about half distance to apex (similar to other Okanopteryx species). Pterostigma broad, 2.5 times longer than wide. Region of possible oblique brace vein missing. Crossveins in postnodal, postsubnodal spaces mostly aligned. Two cells wide distal to pterostigma in region of costal space preserved except one wide at pterostigma. RA–RP1 space one cells wide (apical-most portion not preserved). RP1–IR1 space goes to two cells wide seven cells distal to origin of IR1. IR1: origin two cells distal to origin of RP2; two cells wide close before level of pterostigma. RP2: origin 3.5 cells distal to origin of IR2; two cells wide same level as IR1–RP2 space; linear, straight to slight, even curve posteriorly near margin. IR2 origin at subnodus, mostly straight, linear except near margin slightly zigzag with slight, even curve posteriad. RP3-4: origin about half way between anterior-distal corner of quadrangle, nodus; straight, linear to slight, even curve posteriorly near margin. MA straight, linear, begins slight zigzag shortly beyond level of origin of IR1, increasingly so, but missing close to margin. MP linear, slightly curved from quadrangle, missing near terminus. CuA slightly zigzagged distal to level of origin of RP2; CuA–A space about six columns of three cells wide; otherwise two distal to level of nodus.

Paratype 1, GSC 141101, male ( Figs. 54 , 55A, 55 C–55F). Complete body, total length 37.7 mm; abdomen preserved in ventral aspect, at least distal to segment 4; thorax and head in dorsal aspect; portions of four legs, portions of all four wings. Head ( Fig. 10 L , Table 1 ). In dorsal view about 3.7 mm across eyes at widest point; approximate length from anterior edge of antefrons to posterior of occiput about 1.5 mm; at level of hind ocelli distance between medial margins of eyes about 1.0 mm, width of eye about 1.2 mm, length 1.8 mm. Thorax. Dorsum of prothorax oval, 2.1 mm wide, 1.1 mm long; pterothorax about 6.0 mm long. Legs. Lengths of leg segments unclear, not measured. Wings. Left fore(?)wing. Proximal portion to presumed (based on holotype) half distance between nodus, apex. Darkly infuscate from nodus to distal portion preserved, hyaline proximal to nodus. Maximum width 6.9 mm. Crossveins aligned in postnodal, postsubnodal spaces (portion preserved). RP1 –IR1 space goes to two cells wide three cells distal to origin of IR1. IR1: origin two cells distal to origin of RP2 . RP2 : origin three cells distal to origin of IR2. IR2 origin very close to subnodus; straight, linear. RP3-4 : origin not preserved; linear, straight where reserved. MA linear, slightly curved from quadrangle to zigzag increasingly from about level of origin of IR1. MP linear, slightly curved from quadrangle. CuA with slight zigzag beyond level of nodus, increasing beyond level of origin of RP2 , CuA–A space about 10 columns with 3 cells wide, the rest two distal to level of origin of PR 3-4. Left hind(?) wing. Same region preserved as left fore(?)wing, but more complete in antenodal region. Petiole partially preserved. Shape as for left fore(?)wing. Width 6.9 mm. Hyaline, infuscate in same regions, but infuscation notably lighter. Crossveins in postnodal, postsubnodal spaces aligned. RP1 –IR1 space goes to two cells wide four cells distal to origin of IR1. IR1: origin three cells distal to origin of RP2 . RP2 : origin four cells distal to origin of IR1; straight, linear. IR2 origin very close to subnodus; straight, linear. RP3-4 : origin at level of Ax2, half way between anteriordistal corner of quadrangle, nodus, in middle third between arculus, nodus. MA slightly zigzagged from level of origin of IR1, increasingly so distal to it. MP linear, slightly curved from quadrangle through preserved portion near terminus. CuA linear from level of quadrangle until slight zigzag from nodus level increasingly so after level of origin of RP2 ; CuA–A space with about ten columns of three cells at widest, otherwise two from level of origin of RP3- 4 . Right wings fragmentary, preserved portions do not disagree with left wings; presumed hind(?) wing with same lighter infuscation as left hind(?) wing. Presumed fore(?)wing colouration as in holotype. Abdomen approximately 20.0 mm long. Anal appendages: cerci 1.3 mm long, 0.4 mm wide, evenly curved medially, the rounded apices nearly touching; paraprocts about 0.8 mm long, 0.4 mm wide at base tapering to 0.2 mm at apex, extending about 0.7 the length of cerci; medial side straight, lateral side curved medially; apex slightly hooked medially, rounded .

Paratype 2, F-790 ( Fig. 55 ). Head; thorax, parts of five legs, both forewings missing the apical quarters, the proximal portions of both hind wings; the abdomen to part of segment 3; preserved in dorsal aspect. Head ( Fig. 10 O , Table 1 ). Relatively well preserved; in dorsal view about 3.8 mm across eyes at widest point; approximate length from anterior edge of antefrons to posterior of occiput about 1.7 mm; at level of hind ocelli distance between medial margins of eyes about 1.1 mm, width of eye about 1.3 mm, length 2.2 mm. Thorax. Prothorax roughly oval, about 1.5 mm wide, 1.0 mm long; pterothorax slightly deformed, especially posterior part, about 5.9 mm long, 3.2 mm wide. Legs. Left profemur 3.0 mm long, tibia 3.8 mm (setae 0.6 mm long, their bases separated by about half their length), tarsus 1.5 mm. Right forewing. Petiolate. no accessory antenodal crossveins. Maximum width 6.5 mm. Proximal portion hyaline, darkly infuscate from level of nodus to near level where RP1 –IR1 space becomes two cells wide, hyaline in portion preserved distal to this. Crossveins in postnodal, postsubnodal spaces almost all aligned. RP1 space goes to two cells wide five cells distal to origin of IR1. IR1: origin three cells distal to origin of RP2 . RP2 : origin four cells distal to origin of IR2. IR2: origin at subnodus; linear, straight in portion preserved. RP3-4 : origin about 2/3 from arculus to nodus; linear, straight in portion preserved. MA linear, slightly curved from quadrangle to about level of origin of IR1, increasingly zigzags distal to this. MP linear, slightly curved from quadrangle, small portion near terminus not preserved. CuA increasingly zigzagged from level of origin of RP3-4 ; CuA–A space about seven columns of three cells at widest, otherwise two beyond level of origin of RP3-4 . Left forewing. Petiolate. no accessory antenodal crossveins in areas preserved. Width 6.6 mm. Proximal portion hyaline, darkly infuscate from level of nodus to near level where RP1 –IR1 space becomes two cells wide, hyaline in portion preserved distal to this. Crossveins in postnodal, postsubnodal spaces almost all aligned. RP1 –IR1 space becomes two cells wide four cells distal to origin of IR1. IR1: origin three cells distal to origin of RP2 . RP2 : origin 3.5 cells distal to origin of IR2. IR2: origin at subnodus; linear, straight in portion preserved. RP3-4 : origin similar distance from nodus as right forewing; linear, straight in portion preserved. MA linear, slightly curved from quadrangle to about level of origin of IR1, zigzagged distal to this. MP linear, slightly curved from quadrangle, small portion near terminus not preserved. CuA increasingly zigzagged from level of origin of RP3-4 ; CuA–A space number of columns of three cells wide difficult to determine as preserved, two evident, likely more, otherwise two beyond level of origin of RP3-4 . Right hind wing. Proximal part of wing from articulation to distal side of quadrangle present. Left hind wing. Only base of petiolate portion present. Abdomen mostly missing; only segments, 1, 2 and proximal part of 3 present .

Paratype 3, GSC 141102, two wings ( Fig. 57 ). Middle portions. Wing 1 ( Fig 57B ) missing posterior portion. Darkly infuscate consistent with the holotype. Crossveins in preserved portion of postnodal, postsubnodal spaces aligned. RP1 –IR1 space becomes two cells wide four cells distal to origin of IR1. IR1: origin three cells distal to that of RP2 . RP2 : origin not preserved. Preserved portions of IR1, RP2 , IR2, RP3-4 , MP linear, mostly rather straight. MA becoming, increasingly zigzagged distal to level of origin of IR1. MP linear, slightly curved. Wing 2 ( Fig. 50C ). As for wing 1, except: maximum width 6.8 mm. Crossveins in preserved portion of postnodal, postsubnodal spaces aligned, at most slightly not. RP1 –IR1 space becomes two cells wide three cells distal to origin of IR1. IR1: origin two cells distal to that of RP2 . RP2 : origin 2.5 cells distal to that of IR2. CuA increasingly zigzagged about distal to level of origin of RP2 ;>7 columns of cells three cells deep, the rest two, then one apically .

Paratype 4, RBCM 3087, wing ( Fig. 53 ). Parts of apical half, missing much of apical-most posterior portion. Maximum width 6.9 mm. Proximally-preserved portion infuscate, hyaline apical to this as in holotype. Pterostigma broad, 2.5 times longer than wide. Preserved crossveins in postnodal, postsubnodal spaces aligned or almost so. Two (possibly more?) cells wide distal to pterostigma in costal space (fragmentarily preserved). RP1 –IR1 space becomes two cells wide four cells distal to origin of IR1. Preserved portions of RP2 , IR2, RP3-4 , MP linear, rather straight. MA indistinctly preserved. CuA zigzagged in portion preserved; at least a few columns of three cells wide .

Etymology. The specific epithet is a toponym referring to its only known locality.

Range and age. McAbee, BC, Canada; mid-Ypresian.

Figures

FIGURE 10. Heads of Zygoptera (A, B) and comparative Anisoptera, Anisozygoptera, Tarsophlebiidae, Stenophlebiidae (C– H); and Cephalozygoptera (I–Q).A, Zygoptera, Heteropodagrion superbum Ris redrawn from Fig. 318 of Garrison et al. (2010); B, Zygoptera, Minagrion ribeiroi (Santos) redrawn from Fig. 1842 of Garrison et al. (2010); C, Anisoptera, Gomphidae, Cyanogomphus waltheri Selys redrawn from Fig. 383 of Garrison et al. (2006); D, Anisozygoptera, Epiophlebia laidlawi Tillyard redrawn from Fig. 2 of Büsse (2016); E, Anisoptera, Aeshnidae, Limnetron antarcticum Förster redrawn from Fig. 16 of Garrison et al. (2006); F, Stenophlebia sp. redrawn from Tillyard & Fraser 1940, Fig, 4C; G, Tarsophlebiidae, Tarsophlebia eximia Hagen redrawn from Fleck et al. (2004) Fig. 1C; H, photograph of Tarsophlebia eximia in MCZ collections; I, Sieblosia jucunda redrawn from Hagen (1858, Plate 24, Fig. 1); J, Cephalozygoptera incertae sedis SR 14-003-004; K, Phenacolestes parallelus UCM-4545; L, Okanopteryx macabeensis, GSC 141101; M, Okanagrion beardi paratype 1, RBCM P1546; N, Okanopteryx fraseri GSC 141104; O, Okanopteryx macabeensis F-790; P, Dysagrion fredericii redrawn from Scudder 1890, plate 6, Fig. 10; Q, Dysagrion fredericii redrawn from Scudder 1890, plate 6, Fig. 2. G and H, I, J to O to scales, all 5 mm; others, originals without scales.

FIGURE 50. Okanopteryx fraseri paratype 1, RBCM P1551. A, photograph; B, drawing. Both to scale, 5 mm.

FIGURE 53. Okanopteryx macabeensis holotype, GSC 141100: A, photograph; B, drawing. Paratype 4: RBCM 3087: C, photograph; D, drawing. All to scale, 5 mm.

FIGURE 54. Okanopteryx macabeensis paratype 1, GSC 141101. A, photograph of specimen; B close-up photograph of male anal appendages; C, drawing of male anal appendages labelling abdominal segments 8–10 and indicating paraprocts (red) and cerci (green). A, to 5mm scale, B, C, to 3 mm scale.

FIGURE 55. Okanopteryx macabeensis and O. fraseri. A, O. macabeensis paratype 1, drawing of body; B, holotype O. fraseri drawing of body; abdominal segments labelled; C–F, wings of O. macabeensis paratype 1. A, B and C–F to scales, 5 mm.

FIGURE 56. Okanopteryx macabeensis paratype 2, F-790. A, labelled drawing of wing; B, drawing of specimen; C, photograph of specimen. A to scale, B, C to scale, both scales 5 mm.

FIGURE 57. Okanopteryx macabeensis paratype 3, GSC 141102. A, photograph of specimen; B, drawing of upper wing in A, (vertically inverted for anterior margin on top); C, drawing of bottom wing in A. All to scale, 5 mm.