612:. The suborder was based on the zygopteran body plan, but considered distinct due to the eye to head capsule structure and distances. Unlike zygopterans where the head is 3-5 times as wide as long, in cephalozygopterans the heads are only around double as wide as long making them narrower than in any zygopterans. The distance between the eyes in cephalozygopterans, at about one eye width, was also suggested to be different from known zygopterans where the width is typically at more than doubled. The erection of Cephalozygoptera was criticized by Andre Nel and Darren Zheng (2021) who argued that the noted differences were due in part to deformation of the head and eye areas during deposition and fossilization combined with characters that are present in a larger group of odonates than Archibald, Cannings, and Erickson reported. Based on their review and critique of the justifications for cephalozygoptera, Nel and Zhang considered the name and grouping unwarranted and proposed it to be treated as a synonym of Zygoptera. Archibald and Cannings (2021) responded to the arguments, maintaining that Cephalozygoptera as valid. In subsequent papers both suborders have been used, depending on if the author group includes Archibald or Nel.
1484:(LMA) of the Republic and McAbee paleofloras. The CLAMP results after multiple linear regressions for Republic gave a mean annual temperature of approximately 8.0 °C (46.4 °F), with the LMA giving 9.2 ± 2.0 °C (48.6 ± 3.6 °F). CLAMP results from McAbee returned the higher 10.7 °C (51.3 °F) which was supported by the 10.4 ± 2.4 °C (50.7 ± 4.3 °F) returned from the LMA. These are lower than the mean annual temperature estimates given for the coastal Puget Group, which is estimated to have been between 15–18.6 °C (59.0–65.5 °F). The bioclimatic analysis for Republic and McAbee suggests mean annual precipitation amounts of 115 ± 39 cm (45 ± 15 in) and 108 ± 35 cm (43 ± 14 in) respectively.
963:
853:
115:
813:
1204:
318:
1087:
837:"SR 93-11-02" wasn't identified to either gender or forewing/hindwing in the species description. The wing is 7.0 mm (0.28 in) wide and 25.6 mm (1.01 in) long from arculus to apex, with one diagonal break running across the whole wing, but not distorting the venation. The nodus of the wing is placed 26% of the distance from wing base to apex, and the pterostigma starts 15.3 mm (0.60 in) apically from there. Coloration of the wing is interpreted to be light to hyaline in the basal half transitioning to dark in the apical half. A lighter
128:
874:. The species is the only one to be found in both Washington and British Columbia formations, with the majority of specimens being disarticulated wings, while the only semi-complete body fossil is from McAbee. Of the described species a total of six are from McAbee and housed in British Columbian institutions. The holotype "F-1052", Paratype 5 "F-1140", and Paratype 7 "F-1044" are included in the Thompson Rivers University collections. Paratype 8
651:, Archibald and Cannings treat the assumed hindwings as having a basal hind margin as being smoothly curved from base to level with the quadrangle cell, while the front margin is nearly straight between the nodus and the pterostigma. In the assumed forewings the basal hind margin is mildly convex and widened when reaching the level of the quadrangle cell base, while the front margin between nodus and pterostigma is slightly curved.
499:
1115:
base, in the hind wing the nodus is more apically placed at 30% distance. The pterostigma on both wings are longer than wide, being 3.5 times longer than wide. While the forewing pterostigma is five cells long, however, the hindwing pterostigma is six cells long. Similarly the hindwing CuA–A space is six cells wide, while the forewing CuA–A space is only five cells.
2403:
339:
325:
744:, "RBCM P1546" of the Royal British Columbia Museum collections. Both fossils were recovered from the Hoodoo Face beds, with the collector and collection date of "F-791" unknown, while "RBCM P1546" was collected in June 1998 by an 8th or 9th grade student and donated to the RBCM by Graham Beard. Archibald and Cannings chose the species name "
1378:(2013) showed higher levels of beta diversity in the Eocene Okanagan Highlands, with genera found across several sites, but frequently each site having unique species, a situation that was facilitated by mid-latitude climatic equability during the early Eocene "greenhouse world" which created thermal barriers to species.
1109:
The presumed forewing is wide-oval in outline above the nodus region with the line of symmetry centered on the longitudinal line. The total wing length is 30.5 mm (1.20 in), 27.6 mm (1.09 in) from arculus level to apex, and a length of 16.5 mm (0.65 in) between the nodus
1365:
in the
Ypresian Okanagan region. Local scale diversity, called alpha diversity, was documented in the flora of both sites, and the insects of McAbee, with the diversity level being compared to Costa Rica's modern tropical lowlands. The beta diversity of the Okanagan highlands has also been explored
1230:
The species is most distinct in that the wing membrane is lightly but uniformly darkened, though
Archibald and Cannings note the specimens might appear as hyaline in some fossils. The type series wings are each incomplete, with the holotype missing the basal region up to the nodus. As preserved the
793:
are both indistinct and lacking detail. A 4.1 mm (0.16 in) width across the head at the widest point is reported and the eyes are given 1.5 mm × 2.0 mm (0.059 in × 0.079 in) dimensions. The distance between the eyes is 1.2 mm (0.047 in). The thorax
631:
which is between 3 and 3.5 times wider than tall, with a few exceptions where the width is 4 times wider. The genus is missing the oblique brace vein, which is present or unreported for the majority of dysagrionine species and there are no accessory crossveins immediately basal or apical to the CuP.
1282:
and the latter one by Karl
Volkman. Stonerose specimens "SR 06-08-18", collected by Donald Volkman, and "SR 11-51-07 A&B", collected by Dennis Vickerman are both from the "Boot Hill" site. The Royal British Columbia Museum specimen "RBCM P1550" was collected by John Leahy from Hoodoo Face beds
1217:
is one of the species endemic to
Republic, with three described fossils having been recovered from 1998 to 2016. The holotype, "SR 99-14-02", and paratype 1, "SR 16-006-001", were both recovered from the "Corner lot" while paratype 2, "SR 11-21-09" came from the "Boot hill" site. The earliest found
997:
wing is undetermined, though the wing is noted to be curved above the petiole and rather straight between pterostigma and nodus. The wing has a total length of 30.5 mm (1.20 in), broken down to 25.2 mm (0.99 in) from arculus to apex and 20.3 mm (0.80 in) from the nodus
894:
The general coloration is of the basal 2/5s of the wing is hyaline, while the apical 3/5 is mostly darkened. In the darkened region there is a light to hyaline "u-shaped" window placed so its midpoint is aligned with the midpoint between pterostigma and nodus. The light facia extends almost to the
545:
but did not designate the fossil specimens each identification was based on. The odonate fossil material was studied subsequently by S. Bruce
Archibald and Robert Cannings with a series of papers being published between 2019 and 2021 on the dragonflies and damselflies of the Okanagan Highlands. The
1468:
range between 0.7–1.2 km (0.43–0.75 mi) higher than the coastal forests. This is consistent with the paleoelevation estimates for the lake systems, which range between 1.1–2.9 km (1,100–2,900 m), which is similar to the modern elevation 0.8 km (0.50 mi), but higher.
1114:
hindwings by
Archibald and Cannings. Its a total of 26.5 mm (1.04 in) from arculus to apex, with a nodus to pterostigma distance of 15.7 mm (0.62 in), and a with of 8.5 mm (0.33 in). While in the forewing the nodus is placed about 26% of the distance up from the wing
886:
were all collected by John Leahy before being donated to the Royal
British Columbia Museum collections. The remaining eleven specimens are all housed in the Stonerose Interpretive center collections and donated over a period from 1994 to 2017 from both the "Corner lot" and "Boot Hill" sites. The
802:. The leg sections present are too badly damaged and disarticulated to tell any detail. The abdomen is only present as sections one and two. While the underside of the segment may have remnants of the secondary male genitalia, there isn't enough detail left to confirm the sex of the specimen.
751:
Specimen "F-791" has the better preserved wing venation with one full forewing and portions of at least two other wings, possibly all three present. In addition there are fragments of the abdomen, part of the thorax, and possibly part of one eye preserved as a disarticulated grouping around the
607:
and
Dysagrionidae has been noted to be Zygoptera(?) on occasion. Archibald, Cannings, and Robert Erickson evaluated the known head morphology of the fossils and concluded that they did not belong to any of the three defined odonate suborders, but instead were part of a new subfamily they named
1059:
The wing has a pterostigma to nodus length of 15.4 mm (0.61 in) and a maximum width of 8.5 mm (0.33 in). The wing is missing the apical most area, thus due to the incomplete nature of the wing, a full length estimate was not given and the nodus position was estimated to be
841:
is present in the darkened area between the nodus and pterostigma, extending down from the frontal wing edge to around the wing midpoint. Due to the region not being preserved, the true length of the pterostigma is uncertain, and the width of the costal space apical of the pterostigma is only
1168:
which are fully hyaline and darkened respectively. The wings are hyaline in the basal 2/5ths region and then darkened for the apical 3/5ths. Within the darkened area is a lighter fascia between the nodus and pterostigma, crossing from the front margins to about midway across the wing. The
934:
color patterning in zygopteran species, and thus, having one male present in the group, they opted to take a conservative position on gender of the fossils. They did note that variation in forewing and hindwing coloration is also seen in zygopterans, but they did not consider it present in
1231:
specimen is 22.7 mm (0.89 in) long from nodus to apex, 18.5 mm (0.73 in) long between the pterostigma and nodus, and 7.8 mm (0.31 in) wide. The pterostigma is 3.5 times longer than wide and has a total width of seven cells. The overall wing shape distinguishes
1408:
biotas that are preserved. The highlands temperate biome preserved across a large transect of lakes recorded many of the earliest appearances of modern genera, while also documenting the last stands of ancient lines. The warm temperate highland floras in association with downfaulted
1218:
and most complete, the holotype was discovered on 23 August 1998 by
Providence Worley. The second specimen to be found was paratype 2 donated to Stonerose on 2 August 2008 by Eric Blumhagen, and the most recently found was paratype 1, found on 9 October 2016 by Kattia Rojas. Of the
1413:
and active volcanism are noted to have no exact modern equivalents. This is due to the more seasonally equitable conditions of the Early Eocene, resulting in much lower seasonal temperature shifts. However, the highlands have been compared to the upland ecological islands in the
1047:
as a reference to the amount of dense crossveins found on the wing. Based on the straight front margin between nodus and pterostigma combined with a deeply curving rear margin, the wing is likely a hindwing. The coloration is described by
Archibald and Cannings as "like that of
682:
The specimen is a partial wing indeterminate regarding sex and being forewing or hindwing, as it is missing the basal third of the wing and areas of the apical two-thirds. The wing is overall narrower at the point of the nodus, 5.5 mm (0.22 in) wide, than most other
635:
Between the species, coloration and patterning of the wings are widely varied, most species having broad areas of either light to clear windows on a darkened background, or conversely smaller areas of darkened membrane on light to clear background wing. At least one species
784:
For "RBCM P1546" the wings are mostly absent, with only one well preserved forewing, and a basal section of one hind wing present. However, the head, thorax and the base of the abdomen are all present, along with scraps of the legs. The head is fairly well preserved, though
947:
the forewings and hindwings are articulated, and both have the same color patterning. Conversely, Archibald and Cannings also called out the fluctuation in overall wing size within forewing specimens and hindwing specimens which they considered as a possible result of
577:
veins, while in Dysagrioninae species the placement is around 2/3 of the distance towards the subnodus from the arculus. additionally the IR2 vein in petrolestines branches from very near or on the RP3-4, but forks at or on the subnodus in Dysagrionines. With
1356:
species were likely to be active daytime hunters around the lakes they were eventually entombed in. The genus is noted by Archibald and Cannings for being the most speciose dysagrionid odonate of the highlands, a situation attributed to factors driving both
756:
is 16.8 mm (0.66 in), while the distance from the arculus to pterostigma is 22.0 mm (0.87 in). The overall wing outline on the apical side of the nodus is oval with symmetry centered on the lateral midline. The coloration is similar to
1569:
Archibald, S. B.; Cannings, R. A.; Erickson, R. J.; Bybee, S. M.; Mathewes, R. W. (2021). "The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America".
2363:
Greenwood, D.R.; Archibald, S.B.; Mathewes, R.W; Moss, P.T. (2005). "Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape".
752:
wings. The forewing is 8.3 mm (0.33 in) wide by 31.5 mm (1.24 in) long from the arculus to wing tip, and is only missing a few small areas of apical area and near the nodus. The distance from the nodus to
998:
to apex. At the widest point the oval shaped wing is 8.5 mm (0.33 in) across. The overall arculus to apex length divided by width ratio is distinct, with five species having a greater length to width ratio, and
1226:
has also been found at two Klondike Mountain Formation outcrops. As they did with other species in the genus Archibald and Cannings named the species for Providence Worley in recognition of her contribution to science.
1030:
is known from only the holotype specimen, "RBCM 11799.001" of the Royal British Columbia Museum which was collected from the Hoodoo face beds of McAbee by John Leahy. Archibald and Cannings created the species name
977:
is only known from the holotype wing. The wing was recovered on 29 April 2006 by Gregg Wilson and subsequently donated to the Stonerose Interpretive Center collections as "SR 06-69-17 A&B". The species name
1294:
based on the crossveins in the RA–RP1 space. Due to the poor preservation of "SR 06-08-18" firm placement was considered not possible, but based on the extent of the light facia, it possibly is a specimen of
1440:
climate, in which winter temperatures rarely dropped low enough for snow, and which were seasonably equitable. The paleoforests surrounding the lakes have been described as precursors to the modern
1667:
Moss, PT; Greenwood, DR; Archibald, SB (2005). "Regional and local vegetation community dynamics of the Eocene Okanagan Highlands (British Columbia - Washington State) from palynology".
1765:
Archibald, SB; Bradler, S (2015). "Stem-group stick insects (Phasmatodea) in the early Eocene at McAbee, British Columbia, Canada, and Republic, Washington, United States of America".
722:, the darkened apical area is shorter and the dark mid-wing stripe does not seem to extend fully to the hind margin, but only covers a u-shaped region extending from the front margin.
761:
with the basal area of the wings hyaline, the apical areas darkened, with a light colored to hyaline fascia. While the basal hyaline area also encompasses the basal 2/5 of the wing,
1110:
and pterostigma. The total width is 9.0 mm (0.35 in). The other wing has a more curved rear margin and straighter nodus to pterostigma features that are deemed distinct to
1060:
approximately 28% apical from the wing base. The pterostigma is three times longer than wide, being approximately 5.5 cells long and with oblique basal and apical ends. Unlike
550:
was named by Archibald and Cannings (2021) with a series of eight named species and a group of unplaced fossils which were not placeable to species. They coined the genus name
352:
2242:
Archibald, S.B.; Greenwood, D.R.; Mathewes, R.W. (2013). "Seasonality, montane beta diversity, and Eocene insects: Testing Janzen's dispersal hypothesis in an equable world".
565:
Based on shared characters of the wings, such as a missing crossvein O and the CuA–A space being expanded, Archibald and Cannings placed the new genus into the extinct family
1311:
but none of these species has crossveins as closely spaced as in "UWBM-74307". Lastly "RBCM P1550" is also very poorly preserved, with the closest possible species being
569:. Separation of the two dysagrionid subfamilies is based on the basal origin point of wing vein RP3-4, in the Petrolestinae species this is positioned midway between the
830:
collections as "SR 93-11-02". The specimen was collected and donated by Laurie Dorrell on October 31, 1992, and as such Archibald and Cannings chose to name the species
870:
species, being known from the holotype, a series of nine paratypes, plus a group of seven "additional material" specimens attributed to the species but not included as
1266:
Archibald and Cannings noted that of the Okanagan Highlands fossils that they examined, five specimens from Republic and one fossil from McAbee were identifiable to
317:
2479:
1805:
Archibald, S. B.; Makarkin, V. N. (2021). "Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation".
2142:
Wilf, P.; Cúneo, N.R.; Johnson, K.R.; Hicks, J.F.; Wing, S.L.; Obradovich, J.D. (2003). "High plant diversity in Eocene South America: evidence from Patagonia".
1299:. With "UWBM-74307", the darkening does not extend all the way to the nodus from the apex, eliminating several of the infuscate species. The most similar are
1374:
hypothesis that species in temperature stable environments are less able to pass thermal barriers such as high elevation points. Building on that, Archibald
895:
hind margin in some specimens and all the way across the wing in others. While similarly placed, the window is wider and extends much further than those on
1290:
species. However "SR 11-60-17AB" and "SR 11-51-07" are too fragmentary to suggest species affinity. "SR 94-05-30" is a wing apex which can be excluded from
1405:
1401:
1394:, including the Early Eocene formations between Driftwood Canyon at the north and Republic at the south, have been described as one of the "Great Canadian
930:
Due to the uniformity of the wing coloration, Archibald and Cannings decided to treat the known specimens as all males. This was based on the frequency of
632:
The costal space apical of the pterostigma is between three and four cells wide. The petiole section of the wings ends basally to the CuA and CuP veins.
1102:
was Stonerose Interpretive Center specimen "SR 98-12-10". Archibald and Cannings designated it as the type species of the genus, and named the species
1017:
in having a fully hyaline wing membrane, counter to the other species where light and dark color-patterning is present, or the full wing is darkened.
2466:
597:(damselflys), however the known head morphology of the included species has led to occasional placement questions. The Cretaceous dysagrionid genus
1335:
as male based on the similarity of the wing coloration as a result while noting that not all Zygoptera species have sexually dimorphic coloration.
1270:. Each of the fossils are incomplete, and as such, they can't be placed into the described species, often with several possibilities presented.
2522:
1052:". The basal area is hyaline and the apical area is darkened with a clear to lighter tone fascia extending in a narrower stripe than seen in
2512:
1441:
1429:
The Republic and McAbee upland lake systems were surrounded by a warm temperate ecosystem with nearby volcanism. The highlands likely had a
2527:
2057:"The first Odonata from the early Eocene Allenby Formation of the Okanagan Highlands, British Columbia, Canada (Anisoptera, Aeshnidae and
1160:
Both forewing and hindwing have a very similar color patterning, which is similar to the general coloration of all other species besides
1850:"The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America"
962:
644:
wing. In all species, the fore and hindwings are very similar, so differentiation is difficult. Based on the associated wings of the
2307:"Urticaceae leaves with stinging trichomes were already present in latest early Eocene Okanogan Highlands, British Columbia, Canada"
852:
812:
664:
is one of the four species found at the McAbee Fossil Beds, being known at the time of description from only the holotype fossil,
1703:
2517:
887:
species was named for Shay Hoban who found "paratype 3", SR 94-05-22 A,B, on 15 May 1994, which was the first instance of an
582:
the RP3-4 originates near the 2/3 point towards the subnodus, while the IR2 forks from or directly basal to the subnodus.
1086:
114:
2532:
1632:
Ewing, T.E. (1981). "Regional stratigraphy and structural setting of the Kamloops Group, south-central British Columbia".
827:
665:
395:
291:
1327:
With a limited number of body fossils, the presence of sexually dimorphic wing coloration was not determined. Archibald
463:
is reported from both the "Boot hill" and "Corner lot" sites. Tuffs of the Klondike Mountain Formation had been dated to
838:
786:
574:
570:
1891:"Detrital zircon UPb ages and Hf-isotopes from Eocene intermontane basin deposits of the southern Canadian Cordillera"
1203:
790:
428:
1064:
the major veins are less deeply curved close to the apex where they are preserved. The cells in the apical region of
1169:
coloration differs from other species with patterning in that the light window is narrower or shorter in depth. In
127:
737:
1387:
365:
271:
1613:
Hills, L.V.; Baadsgaard, H. (1967). "Potassium-argon dating of some Lower Tertiary strata in British Columbia".
436:
2101:
1177:
the light window is three times wider and extends nearly to or all the way to the hind margin. The width of
669:
593:
is controversial. Until 2021 the family had been treated as likely belonging to the living odonate suborder
2421:
1971:
Nel, A.; Zheng, D. (2021). "The recently proposed odonatan 'suborder' Cephalozygoptera: fact or fiction".
1940:"Fossil dragonflies (Odonata: Anisoptera) from the early Eocene Okanagan Highlands, western North America"
387:
2407:
623:
species are united by a series of features that are not seen in total in any other Dysagrioninae taxon.
826:
is known from the lone holotype fossil collected from the Klondike Mountain Formation and housed in the
399:
748:" in recognition of Beard for his long dedication and energy towards British Columbian paleontology.
609:
2453:
2373:
2251:
2151:
1902:
1715:
1676:
1641:
1386:
Both the Republic and McAbee sites are part of a larger fossil site system collectively known as the
1106:
in recognition of finder Carolyn Threadgill. The specimen has both a presumed forewing and hindwing.
432:
283:
1193:
is similar in extent across, the other two have windows that extend much closer to the rear margin.
993:
As with many of the other species known only from disarticulated wings, the sex and position of the
1457:
590:
566:
338:
264:
205:
192:
486:, for which multiple fossils have been recovered from both Republic sites and at McAbee, the only
2336:
2286:
1. Early Eocene Lagerstätten of the Okanagan Highlands (British Columbia and Washington State)".
2224:
2175:
2124:
2082:
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1988:
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1830:
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1595:
1344:
931:
414:
391:
287:
279:
122:
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to be the dominant odonates of the Eocene Okanagan highlands forests. As with modern odonates,
1129:
by the positioning of the MP and CuA veins when they terminate on the apical margin region. For
599:
2484:
511:
The first brief report of an Okanagan Highlands fossil odonate was by Standley Lewis in a 1992
2328:
2216:
2167:
2029:
1871:
1822:
1587:
1423:
1415:
949:
534:
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2381:
2318:
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Archibald, S.; Greenwood, D.; Smith, R.; Mathewes, R.; Basinger, J. (2011). "Great Canadian
2259:
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2019:
1980:
1951:
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1814:
1774:
1723:
1684:
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wing has an IR2 vein which forked from the RP1-2 vein origin, a fork point only shared with
406:
in the 1960s based on ash samples exposed in the lake bed. These samples yielded an age of ~
295:
2305:
DeVore, M. L.; Nyandwi, A.; Eckardt, W.; Bizuru, E.; Mujawamariya, M.; Pigg, K. B. (2020).
1889:
Rubino, E.; Leier, A.; Cassel, E.; Archibald, S.; Foster-Baril, Z.; Barbeau, D. Jr (2021).
1278:
and "SR 11-60-17 A&B" were all collected from the "Corner Lot" site, the former two by
668:
specimen "RBCM.EH2017.050.11.2491" found by John Leahy. Archibald and Cannings coined the
302:
resulting from climatic equitability during the Early Eocene in combination with resultant
298:, where four species are present. The species richness is attributed to high latitude high
1848:
Archibald, S. B.; Cannings, R. A.; Erickson, R. J.; Bybee, S. M.; Mathewes, R. W. (2021).
1461:
1358:
538:
503:
299:
1396:
2377:
2255:
2155:
1906:
1719:
1680:
1645:
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All the specimens from Republic sites are identifiable at least to any of the infuscate
777:
the fascia extends across the will wing width, while it doesn't fully cross the wing in
2195:
1419:
1362:
1036:
604:
559:
303:
2501:
2340:
2128:
2086:
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1992:
1924:
1834:
1599:
1445:
1430:
871:
324:
267:. The genus was first described in 2021 with a series of eight species included from
72:
2228:
2179:
1984:
1786:
1740:
1735:
17:
2444:
1433:
1271:
915:
is the most similar, but the holotype differs in venation in the preserved areas.
498:
268:
2263:
1915:
1890:
521:
from Republic. Four years later a list of insects from Republic was published by
1453:
1437:
1279:
1010:
but they differ on a number of vein and cell characters, as well as coloration.
795:
753:
628:
522:
403:
47:
2024:
2007:
695:
at 4.5 mm (0.18 in) and 4.0 mm (0.16 in) respectively. The
470:, the youngest of the Okanagan Highlands sites, though a revised oldest age of
1866:
1849:
1818:
1583:
799:
794:
has an overall dark coloration, with a possible light stripe running over the
703:, while the other species have fork points between the RP1-2 and RP3-4 veins.
530:
517:
92:
57:
2220:
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260:
159:
139:
97:
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2120:
2033:
1875:
1826:
1591:
2402:
1157:, with arculus through apex/width ratios being 2.6 and 3.1 respectively.
982:
was created by Archibald and Cannings as a combination of the Latin words
2438:
1391:
741:
648:
586:
542:
275:
87:
82:
67:
62:
52:
2471:
2077:
2056:
1956:
1939:
1704:"Seasonality, the latitudinal gradient of diversity, and Eocene insects"
1243:. At 4.5 mm (0.18 in) it is narrower in the nodus region than
1778:
1702:
Archibald, S.B.; Bossert, W.H.; Greenwood, D.R.; Farrell, B.D. (2010).
1473:
641:
555:
465:
256:
179:
102:
77:
2323:
2306:
1098:
Found at the "Corner Lot" site A0307 on 8 April 1998, the holotype of
515:
article where he illustrated two fossils tentatively identified as (?)
472:
419:
1449:
1247:
which is 5.5 mm (0.22 in). The mostly straight MP vein of
398:
in central British Columbia. The lake sediments at McAbee were first
169:
149:
2415:
2385:
1727:
1688:
1653:
923:, where the wing is lightly colored across the entire membrane, and
558:
honoring the Okanagan Highlands plus the suffix "-agrion", from the
408:
368:
of Washington and British Columbia. Of the eight described species,
2212:
2102:"The odonatan insects from the Paleocene of Menat, central France"
1202:
1085:
961:
851:
811:
679:
meaning "narrow" in reference to the overall wing shape estimate.
673:
497:
427:
Four additional species are known exclusively from fossils of the
2458:
274:
sites in western North America. The genus is known from the Late
457:
is only known from the "Corner lot" site A0307 and one species,
2419:
1444:
of Eastern North America and Eastern Asia. Based on the fossil
1013:
The species is also distinguishable from all other members of
911:, all of which are between 1/3 to 1/2 as wide. Coloration of
1400:" based on the diversity, quality and unique nature of the
736:
is known from two specimens, the holotype, "F-791", in the
451:
are known only from the "Boot Hill" site UWBM B4131, while
1319:
based on the CuA–A space having either five or six cells.
529:
article. They expanded the tentative families to include
715:
in having a longer length of the Ax1 vein to the nodus.
306:
between sites due to impassible topographical barriers.
1564:
1562:
1560:
1558:
1556:
1554:
1552:
1550:
1548:
1546:
1544:
1542:
1540:
1538:
1536:
1534:
1532:
1530:
1528:
1526:
1524:
1522:
1520:
1518:
1516:
1189:
are all closer in width, though still wider, and while
769:
in the size of the fascia which is only 1/2 as wide in
687:
specie. Only two species have a narrower nodus point,
1514:
1512:
1510:
1508:
1506:
1504:
1502:
1500:
1498:
1496:
1255:
where the vein has a zig-zagged path near the margin.
927:
where the wing is hyaline across the entire membrane.
477:
was given based on isotopic data published in 2021.
364:
fossils have been found at two sites belonging to the
1141:
the space is widening. The wings are narrower than
286:
where five species are present, and from the coeval
2428:
1133:the two are placed subparallel to each other while
562:"ἄγριος", commonly used in damselfly genus names.
2194:
1739:
1331:opted to consider all 17 illustrated specimens of
2244:Palaeogeography, Palaeoclimatology, Palaeoecology
1153:has a smaller length to width ratio than that of
2358:
2356:
2354:
2352:
2350:
2196:"Why Mountain Passes are Higher in the Tropics"
1938:S. Bruce Archibald; Robert A. Cannings (2019).
1800:
1798:
1796:
1760:
1758:
413:; however, dating published in 2005 provided a
1460:of Western Washington, which are described as
1274:specimen "UWBM-74307" and Stonerose specimens
973:One of the "Boot Hill" site Republic species,
1372:Why mountain passes are higher in the tropics
468: ± 0.54 million years ago
439:, northeast Central Washington. Two species,
422: ± 0.83 million years ago
8:
490:species to be found in multiple formations.
475: ± 0.1 million years ago
417:radiometric date placing the McAbee site at
2416:
2277:
2275:
2273:
2055:Archibald, S. B.; Cannings, R. A. (2022).
2006:Archibald, S. B.; Cannings, R. A. (2021).
1478:climate leaf analysis multivariate program
1448:the lakes were higher and cooler than the
1068:are more densely packed then in any other
113:
31:
2322:
2076:
2023:
1955:
1914:
1865:
2008:"The head of Cephalozygoptera (Odonata)"
1056:most of the way across the wing width.
1492:
278:sediments exposed in northeast central
1615:Canadian Petroleum Geologists Bulletin
1137:has a wide space between them, and in
603:has had affiliation with the suborder
1442:temperate broadleaf and mixed forests
7:
866:is the most common of the described
773:. Additionally in some specimens of
507:with nodus and arculus veins labeled
718:While the coloration is similar to
480:The widest distribution is that of
2366:Canadian Journal of Earth Sciences
1669:Canadian Journal of Earth Sciences
1634:Canadian Journal of Earth Sciences
25:
2061:Cephalozygoptera, Dysagrionidae)"
1452:coastal forests preserved in the
2401:
891:fossil being found and donated.
337:
323:
316:
126:
1985:10.11646/palaeoentomology.4.2.5
1235:from all other species except
919:is most easily separated from
1:
2523:Fossil taxa described in 2021
2109:Acta Palaeontologica Polonica
2100:Nel, A.; Jouault, C. (2022).
1464:ecosystems. Estimates of the
1121:wings are distinguished from
828:Stonerose Interpretive Center
666:Royal British Columbia Museum
525:and Lisa Barksdale in a 1996
2513:Eocene life of North America
2264:10.1016/j.palaeo.2012.10.043
1916:10.1016/j.sedgeo.2021.105969
27:Extinct genus of damselflies
2528:Klondike Mountain Formation
1002:being narrower. In outline
842:estimated as 3 cells wide.
429:Klondike Mountain Formation
2549:
2311:American Journal of Botany
2193:Janzen, Daniel H. (1967).
2025:10.11646/zootaxa.5047.1.10
738:Thompson Rivers University
541:or Megapodagrionidae, and
494:History and classification
2065:The Canadian Entomologist
1944:The Canadian Entomologist
1867:10.11646/zootaxa.4934.1.1
1819:10.11646/zootaxa.4951.1.2
1767:The Canadian Entomologist
1584:10.11646/zootaxa.4934.1.1
1388:Eocene Okanagan Highlands
640:is noted to have a fully
366:Eocene Okanagan Highlands
239:
234:
123:Scientific classification
121:
112:
34:
1348:were noted by Archibald
1094:illustration of venation
975:Okanagrion liquetoalatum
967:Okanagrion liquetoalatum
957:Okanagrion liquetoalatum
882:and additional material
2201:The American Naturalist
2164:10.1126/science.1080475
1860:(1): zootaxa.4934.1.1.
1578:(1): zootaxa.4934.1.1.
1476:have been derived from
1474:mean annual temperature
1462:lowland tropical forest
1100:Okanagrion threadgillae
1091:Okanagrion threadgillae
1081:Okanagrion threadgillae
1039:"Λόχμη", anglicized to
411: million years ago
2121:10.4202/app.00960.2021
1251:distinguishes it from
1211:
1095:
970:
860:
820:
554:as a combination of a
508:
386:are only found at the
2518:Fossil Odonata genera
1206:
1089:
965:
939:due to "paratype 7" #
855:
815:
627:species wings have a
501:
400:radiometrically dated
2533:Tranquille Formation
2412:at Wikimedia Commons
1482:leaf margin analysis
824:Okanagrion dorrellae
817:Okanagrion dorrellae
807:Okanagrion dorrellae
740:collection, and the
442:Okanagrion dorrellae
388:Tranquille Formation
353:class=notpageimage|
18:Okanagrion dorrellae
2378:2005CaJES..42..167G
2256:2013PPP...371....1A
2156:2003Sci...300..122W
2078:10.4039/tce.2022.16
1957:10.4039/tce.2019.61
1907:2021SedG..42205969R
1895:Sedimentary Geology
1720:2010Pbio...36..374A
1681:2005CaJES..42..187M
1646:1981CaJES..18.1464E
1458:Chuckanut Formation
1424:African rift valley
1215:Okanagrion worleyae
1198:Okanagrion worleyae
1006:is most similar to
986:meaning clear, and
732:The McAbee species
662:Okanagrion angustum
656:Okanagrion angustum
356:McAbee and Republic
1901:: Article 105969.
1779:10.4039/tce.2015.2
1434:upper microthermal
1212:
1096:
1028:Okanagrion lochmum
1022:Okanagrion lochmum
990:meaning "winged".
971:
932:sexually dimorphic
861:
821:
707:also differs from
527:Washington Geology
513:Washington Geology
509:
392:McAbee Fossil Beds
288:McAbee Fossil Beds
272:Okanagan Highlands
2495:
2494:
2422:Taxon identifiers
2406:Media related to
2324:10.1002/ajb2.1548
2317:(10): 1449–1456.
2288:Geoscience Canada
2150:(5616): 122–125.
1472:Estimates of the
1438:lower mesothermal
1416:Virunga Mountains
1411:lacustrine basins
950:sexual dimorphism
864:Okanagrion hobani
857:Okanagrion hobani
847:Okanagrion hobani
734:Okanagrion beardi
727:Okanagrion beardi
535:Megapodagrionidae
483:Okanagrion hobani
248:
247:
230:
43:Ypresian–Ypresian
16:(Redirected from
2540:
2488:
2487:
2475:
2474:
2462:
2461:
2449:
2448:
2447:
2417:
2405:
2390:
2389:
2360:
2345:
2344:
2326:
2302:
2296:
2295:
2279:
2268:
2267:
2239:
2233:
2232:
2207:(919): 233–249.
2198:
2190:
2184:
2183:
2139:
2133:
2132:
2106:
2097:
2091:
2090:
2080:
2052:
2046:
2045:
2027:
2003:
1997:
1996:
1973:Palaeoentomology
1968:
1962:
1961:
1959:
1935:
1929:
1928:
1918:
1886:
1880:
1879:
1869:
1845:
1839:
1838:
1802:
1791:
1790:
1762:
1753:
1752:
1750:
1749:
1743:
1738:. Archived from
1699:
1693:
1692:
1664:
1658:
1657:
1640:(9): 1464–1477.
1629:
1623:
1622:
1610:
1604:
1603:
1566:
1382:Paleoenvironment
1368:Daniel H. Janzen
1342:and the related
1162:O. liquetoalatum
1151:O. liquetoalatum
1072:species besides
1004:O. liquetoalatum
995:O. liquetoalatum
925:O. liquetoalatum
834:in recognition.
670:specific epithet
638:O. liquetoalatum
610:Cephalozygoptera
476:
469:
448:O. liquetoalatum
423:
412:
341:
327:
320:
296:British Columbia
224:
217:
204:
191:
131:
130:
117:
107:
44:
40:Temporal range:
32:
21:
2548:
2547:
2543:
2542:
2541:
2539:
2538:
2537:
2498:
2497:
2496:
2491:
2483:
2478:
2470:
2465:
2457:
2452:
2443:
2442:
2437:
2424:
2398:
2393:
2386:10.1139/e04-100
2362:
2361:
2348:
2304:
2303:
2299:
2281:
2280:
2271:
2241:
2240:
2236:
2192:
2191:
2187:
2141:
2140:
2136:
2104:
2099:
2098:
2094:
2054:
2053:
2049:
2005:
2004:
2000:
1970:
1969:
1965:
1937:
1936:
1932:
1888:
1887:
1883:
1847:
1846:
1842:
1804:
1803:
1794:
1764:
1763:
1756:
1747:
1745:
1728:10.1666/09021.1
1701:
1700:
1696:
1689:10.1139/E04-095
1666:
1665:
1661:
1654:10.1139/e81-137
1631:
1630:
1626:
1612:
1611:
1607:
1568:
1567:
1494:
1490:
1384:
1325:
1297:O. threadgillae
1264:
1201:
1155:O. threadgillae
1149:. The wing of
1145:but wider than
1131:O. threadgillae
1119:O. threadgillae
1093:
1084:
1025:
960:
909:O. threadgillae
850:
810:
730:
659:
646:O. threadgillae
618:
539:Platycnemididae
504:Ischnura aurora
496:
471:
464:
454:O. threadgillae
418:
407:
359:
358:
357:
355:
349:
348:
347:
346:
342:
334:
333:
332:
328:
312:
300:alpha diversity
223:
215:
202:
189:
125:
108:
106:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
42:
41:
38:
28:
23:
22:
15:
12:
11:
5:
2546:
2544:
2536:
2535:
2530:
2525:
2520:
2515:
2510:
2500:
2499:
2493:
2492:
2490:
2489:
2476:
2463:
2450:
2434:
2432:
2426:
2425:
2420:
2414:
2413:
2397:
2396:External links
2394:
2392:
2391:
2372:(2): 167–185.
2346:
2297:
2269:
2234:
2213:10.1086/282487
2185:
2134:
2115:(3): 631–648.
2092:
2047:
1998:
1979:(2): 165–170.
1963:
1950:(6): 783–816.
1930:
1881:
1840:
1792:
1773:(6): 744–753.
1754:
1714:(3): 374–398.
1694:
1675:(2): 187–204.
1659:
1624:
1605:
1491:
1489:
1486:
1466:paleoelevation
1420:Albertine Rift
1383:
1380:
1363:beta diversity
1324:
1321:
1263:
1260:Incertae sedis
1257:
1222:species, only
1200:
1195:
1083:
1078:
1024:
1019:
959:
954:
872:type specimens
849:
844:
809:
804:
729:
724:
658:
653:
617:
614:
605:Anisozygoptera
495:
492:
351:
350:
344:
343:
336:
335:
330:
329:
322:
321:
315:
314:
313:
311:
308:
304:beta diversity
255:is an extinct
246:
245:
244:
243:
237:
236:
232:
231:
213:
209:
208:
200:
196:
195:
187:
183:
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61:
56:
51:
46:
45:
39:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
2545:
2534:
2531:
2529:
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2524:
2521:
2519:
2516:
2514:
2511:
2509:
2506:
2505:
2503:
2486:
2481:
2477:
2473:
2468:
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2460:
2455:
2451:
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2440:
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2342:
2338:
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2330:
2325:
2320:
2316:
2312:
2308:
2301:
2298:
2294:(4): 155–164.
2293:
2289:
2285:
2278:
2276:
2274:
2270:
2265:
2261:
2257:
2253:
2249:
2245:
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2222:
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2214:
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2161:
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2110:
2103:
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2070:
2066:
2062:
2060:
2051:
2048:
2043:
2039:
2035:
2031:
2026:
2021:
2018:(1): 97–100.
2017:
2013:
2009:
2002:
1999:
1994:
1990:
1986:
1982:
1978:
1974:
1967:
1964:
1958:
1953:
1949:
1945:
1941:
1934:
1931:
1926:
1922:
1917:
1912:
1908:
1904:
1900:
1896:
1892:
1885:
1882:
1877:
1873:
1868:
1863:
1859:
1855:
1851:
1844:
1841:
1836:
1832:
1828:
1824:
1820:
1816:
1812:
1808:
1801:
1799:
1797:
1793:
1788:
1784:
1780:
1776:
1772:
1768:
1761:
1759:
1755:
1744:on 2011-08-07
1742:
1737:
1733:
1729:
1725:
1721:
1717:
1713:
1709:
1705:
1698:
1695:
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1678:
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1407:
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1269:
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1250:
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1238:
1234:
1228:
1225:
1221:
1216:
1209:
1205:
1199:
1196:
1194:
1192:
1188:
1184:
1180:
1176:
1172:
1167:
1163:
1158:
1156:
1152:
1148:
1144:
1140:
1136:
1132:
1128:
1124:
1120:
1116:
1113:
1107:
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1101:
1092:
1088:
1082:
1079:
1077:
1075:
1071:
1067:
1063:
1057:
1055:
1051:
1046:
1042:
1038:
1034:
1029:
1023:
1020:
1018:
1016:
1011:
1009:
1005:
1001:
996:
991:
989:
985:
981:
980:liquetoalatum
976:
968:
964:
958:
955:
953:
951:
946:
942:
938:
933:
928:
926:
922:
918:
914:
910:
906:
902:
898:
892:
890:
885:
881:
878:, Paratype 9
877:
873:
869:
865:
858:
854:
848:
845:
843:
840:
835:
833:
829:
825:
818:
814:
808:
805:
803:
801:
797:
792:
788:
782:
780:
776:
772:
768:
765:differs from
764:
760:
755:
749:
747:
743:
739:
735:
728:
725:
723:
721:
716:
714:
710:
706:
702:
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690:
686:
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678:
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671:
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663:
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652:
650:
647:
643:
639:
633:
630:
626:
622:
615:
613:
611:
606:
602:
601:
596:
592:
591:Dysagrionidae
589:placement of
588:
583:
581:
576:
572:
568:
567:Dysagrionidae
563:
561:
557:
553:
549:
544:
540:
536:
532:
528:
524:
520:
519:
514:
506:
505:
500:
493:
491:
489:
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265:Dysagrionidae
263:-like family
262:
259:genus in the
258:
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206:Dysagrioninae
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30:
19:
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2284:Lagerstätten
2283:
2247:
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2237:
2204:
2200:
2188:
2147:
2143:
2137:
2112:
2108:
2095:
2068:
2064:
2058:
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2015:
2011:
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1976:
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1966:
1947:
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1933:
1898:
1894:
1884:
1857:
1853:
1843:
1813:(1): 41–79.
1810:
1806:
1770:
1766:
1746:. Retrieved
1741:the original
1711:
1708:Paleobiology
1707:
1697:
1672:
1668:
1662:
1637:
1633:
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1480:(CLAMP) and
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1397:Lagerstätten
1395:
1385:
1375:
1371:
1353:
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1343:
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1337:
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1328:
1326:
1323:Paleobiology
1316:
1312:
1309:O. dorrellae
1308:
1304:
1300:
1296:
1291:
1287:
1285:
1275:
1272:Burke Museum
1267:
1265:
1259:
1253:O. dorrellae
1252:
1248:
1244:
1241:O. dorrellae
1240:
1236:
1232:
1229:
1223:
1219:
1214:
1213:
1207:
1197:
1191:O. dorrellae
1190:
1186:
1183:O. dorrellae
1182:
1178:
1174:
1170:
1165:
1161:
1159:
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1143:O. dorrellae
1142:
1139:O. dorrellae
1138:
1134:
1130:
1127:O. dorrellae
1126:
1122:
1118:
1117:
1111:
1108:
1104:threadgillae
1103:
1099:
1097:
1090:
1080:
1073:
1069:
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1061:
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1053:
1049:
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1043:which means
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904:
901:O. dorrellae
900:
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893:
888:
883:
879:
875:
867:
863:
862:
856:
846:
836:
831:
823:
822:
816:
806:
798:area on the
783:
778:
774:
770:
766:
762:
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745:
733:
731:
726:
719:
717:
713:O. dorrellae
712:
708:
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696:
693:O. dorrellae
692:
688:
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584:
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441:
440:
437:Ferry County
426:
382:
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375:
370:
369:
361:
360:
310:Distribution
269:early Eocene
251:
250:
249:
226:
219:
218:
35:
29:
1454:Puget Group
1418:within the
1406:paleofaunal
1402:paleofloral
1345:Okanopteryx
1338:Species of
1301:O. angustum
1283:at McAbee.
1280:Wesley Wehr
1276:SR 94-05-30
1249:O. worleyae
1245:O. angustum
1237:O. angustum
1233:O. worleyae
1208:O. worleyae
1171:O. angustum
1166:O. worleyae
1147:O. angustum
1112:Okanagrion
1074:O. angustum
1035:" from the
1000:O. angustum
921:O. worleyae
913:O. angustum
796:mesopleural
754:pterostigma
705:O. angustum
697:O. angustum
689:O. worleyae
629:pterostigma
616:Description
523:Wesley Wehr
460:O. worleyae
431:exposed at
404:K-Ar method
396:Cache Creek
371:O. angustum
294:in Central
292:Cache Creek
199:Subfamily:
2502:Categories
2445:Q124831889
2430:Okanagrion
2409:Okanagrion
2071:(1): e29.
1748:2021-11-13
1621:: 138–149.
1488:References
1354:Okanagrion
1340:Okanagrion
1317:O. lochmum
1292:O. lochmum
1288:Okanagrion
1268:Okanagrion
1220:Okanagrion
1187:O. lochmum
1070:Okanagrion
1066:O. lochmum
1015:Okanagrion
905:O. lochmum
884:RBCM P1549
880:RBCM P1548
876:RBCM P1547
868:Okanagrion
800:mesothorax
685:Okanagrion
625:Okanagrion
621:Okanagrion
600:Congqingia
587:subordinal
580:Okanagrion
552:Okanagrion
548:Okanagrion
546:new genus
531:Euphaeidae
518:Zacallites
488:Okanagrion
402:using the
383:O. lochmum
362:Okanagrion
280:Washington
252:Okanagrion
225:Archibald
220:Okanagrion
160:Arthropoda
36:Okanagrion
2341:225050834
2221:0003-0147
2129:249299630
2087:250035713
2042:244256746
1993:235536486
1925:237717862
1835:233411745
1600:232337536
1392:highlands
1366:based on
1333:O. hobani
1313:O. hobani
1305:O. beardi
1224:O. hobani
1179:O. beardi
1175:O. hobani
1135:O. beardi
1123:O. beardi
1062:O. hobani
1054:O. hobani
1050:O. beardi
1008:O. hobani
937:O. hobani
917:O. hobani
897:O. beardi
889:O. hobani
832:dorrellae
779:O. beardi
775:O. hobani
771:O. beardi
767:O. hobani
763:O. beardi
759:O. hobani
720:O. hobani
709:O. beardi
701:O. beardi
672:from the
595:Zygoptera
377:O. beardi
261:damselfly
242:8 species
146:Kingdom:
140:Eukaryota
2485:11919673
2472:11163249
2459:61249259
2439:Wikidata
2333:33091153
2229:84408071
2180:20101200
2172:12677065
2034:34811004
1876:33756770
1827:33903413
1787:86608533
1736:55208851
1592:33756770
1210:holotype
969:holotype
859:paratype
819:holotype
742:paratype
677:angustum
649:holotype
575:subnodus
543:Lestidae
433:Republic
394:west of
345:Republic
284:Republic
276:Ypresian
235:Species
186:Family:
156:Phylum:
150:Animalia
136:Domain:
2508:Odonata
2374:Bibcode
2252:Bibcode
2250:: 1–8.
2152:Bibcode
2144:Science
2012:Zootaxa
1903:Bibcode
1854:Zootaxa
1807:Zootaxa
1716:Bibcode
1677:Bibcode
1642:Bibcode
1572:Zootaxa
1422:of the
1390:. The
1262:fossils
1045:thicket
1033:lochmum
791:clypeus
642:hyaline
571:arculus
556:toponym
257:odonate
212:Genus:
180:Odonata
176:Order:
170:Insecta
166:Class:
2339:
2331:
2227:
2219:
2178:
2170:
2127:
2085:
2040:
2032:
1991:
1923:
1874:
1833:
1825:
1785:
1734:
1598:
1590:
1450:coeval
1446:biotas
1307:, and
1185:, and
1041:lóchmi
988:alatus
984:liquet
945:F-1044
941:F-1044
839:fascia
787:labrum
746:beardi
380:, and
331:McAbee
229:, 2021
227:et al.
2480:IRMNG
2337:S2CID
2225:S2CID
2176:S2CID
2125:S2CID
2105:(PDF)
2083:S2CID
2038:S2CID
1989:S2CID
1921:S2CID
1831:S2CID
1783:S2CID
1732:S2CID
1596:S2CID
1431:mesic
1376:et al
1359:alpha
1350:et al
1329:et al
1037:Greek
943:. In
674:Latin
560:Greek
466:49.42
415:Ar-Ar
290:near
2467:GBIF
2329:PMID
2217:ISSN
2168:PMID
2030:PMID
2016:5047
1872:PMID
1858:4934
1823:PMID
1811:4951
1588:PMID
1576:4934
1456:and
1404:and
1361:and
1315:and
1239:and
1173:and
1164:and
1125:and
789:and
711:and
691:and
585:The
573:and
473:51.2
445:and
420:52.9
48:PreꞒ
2454:EoL
2382:doi
2319:doi
2315:107
2260:doi
2248:371
2209:doi
2205:101
2160:doi
2148:300
2117:doi
2073:doi
2069:154
2059:cf.
2020:doi
1981:doi
1952:doi
1948:151
1911:doi
1899:422
1862:doi
1815:doi
1775:doi
1771:147
1724:doi
1685:doi
1650:doi
1580:doi
1436:to
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435:in
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