New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks

The osmylid subfamily Protosmylinae is revised based on our emended diagnosis: Petrushevskia Martynova, 1958 and Mesosmylidus Jepson et al., 2012 are excluded (both considered Osmylidae incertae sedis), and Sogjuta Martynova, 1958 is transferred to it from the Mesosmylininae. The late Eocene genus Protosmylus Krüger, 1913 is considered a junior synonym of Osmylidia Cockerell, 1908 based on a distinct apomorphy (deeply forked MA in the hind wing), syn. nov. Three new species of Osmylidia from the early Eocene of North America are described: O. donnae sp. nov. from Quilchena, O. glastrai sp. nov. from Republic, Washington, USA, and an unnamed species of Osmylidia is reported from Driftwood Canyon Provincial Park, British Columbia (all localities of the Okanagan Highlands series), and O. taliae sp. nov. from the Green River Formation of Colorado, USA.

A new genus and species of Euphaeidae (Odonata, Zygoptera) from the early Eocene Okanagan Highlands locality at Republic, Washington, U.S.A.

We describe Republica weatbrooki, a new genus and species of damselfly (Odonata, Zygoptera, Euphaeidae, Eodichromatinae) from the early Eocene (Ypresian) fossil locality at Republic, Washington, U.S.A. Its single specimen is the sole damselfly known from the Okanagan Highlands series of localities in far-western North America.

Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation

Four new species of Raphidiidae are described from the early Eocene of western North America: Megaraphidia antiquissima sp. nov. from McAbee, M. ootsa sp. nov. from Driftwood Canyon, M. hopkinsi sp. nov. from the Allenby Formation (all from British Columbia, Canada), M. klondika sp. nov. from Republic (Washington, United States of America). Archiinocellia Handlirsch, 1910, Archiinocellia oligoneura Handlirsch, 1910 from Horsefly River (British Columbia, Canada), and A. protomaculata (Engel, 2011), comb. nov., from the Green River Formation (Colorado, United States of America) are redescribed. Archiinocellia is assigned to Raphidiidae, sit. nov. The apparent absence of sclerotized gonocoxites 9 in the Archiinocellia protomaculata male is probably plesiomorphic at the family level. As some modern snakeflies do not require a cold interval to complete their development and Eocene Inocelliidae and Raphidiidae lived in regions of warm winters (especially A. protomaculata), adaptation to cold winters in many modern snakeflies is a post-Eocene phenomenon. Eocene Raphidiidae of Europe (Priabonian) differ greatly from those of North America (Ypresian and Priabonian). This pattern might reflect dispersal in either direction or ranges established prior to continental separation. Eocene Inocelliidae of Europe (Priabonian), however, are more similar to those of North America (Ypresian and Lutetian).

The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America

We describe the Cephalozygoptera, a new, extinct suborder of Odonata, composed of the families Dysagrionidae and Sieblosiidae, previously assigned to the Zygoptera, and possibly the Whetwhetaksidae n. fam. The Cephalozygoptera is close to the Zygoptera, but differs most notably by distinctive head morphology. It includes 59 to 64 species in at least 19 genera and one genus-level parataxon. One species is known from the Early Cretaceous (Congqingia rhora Zhang), possibly three from the Paleocene, and the rest from the early Eocene through late Miocene. We describe new taxa from the Ypresian Okanagan Highlands of British Columbia, Canada and Washington, United States of America: 16 new species of Dysagrionidae of the existing genus Dysagrion (D. pruettae); the new genera Okanagrion (O. threadgillae, O. hobani, O. beardi, O. lochmum, O. angustum, O. dorrellae, O. liquetoalatum, O. worleyae, all new species); Okanopteryx (O. jeppesenorum, O. fraseri, O. macabeensis, all new species); Stenodiafanus (S. westersidei, new species); the new genus-level parataxon Dysagrionites (D. delinei new species, D. sp. A, D. sp. B, both new); and one new genus and species of the new family Whetwhetaksidae (Whetwhetaksa millerae).

A new genus and species of split-footed lacewings (Neuroptera) from the early Eocene of western Canada and revision of the subfamily affinities of Mesozoic Nymphidae

A new genus and new species of Nymphidae (Neuroptera) is described from the Ypresian Okanagan Highlands locality of Falkland, British Columbia, Canada: Epinesydrion falklandensisnew genus, new species. This is only the fourth known Cenozoic adult specimen, and all others are less complete. It is the second specimen from the Okanagan Highlands. Currently Nymphidae has two recognised subfamilies. All Cenozoic fossils are confident members of the Nymphinae, but the subfamily assignments of almost all Mesozoic genera are problematic. The Late Cretaceous Dactylomyius is the only genus that might belong to Myiodactylinae. The rest may belong to the undefined stem groups of the family or to the Nymphinae, with varying levels of probability. Mesonymphes sibirica is transferred to Nymphites Haase: N. sibiricus (Ponomarenko), new combination; Sialium minor to Spilonymphes Shi, Winterton, and Ren: Spilonymphes minor (Shi, Winterton, and Ren), new combination; “Mesonymphes” apicalis does not belong to Mesonymphes Carpenter and may not even belong to the Nymphidae. The fossil record of the family occurs across much of the globe, but today they are restricted to Australia, New Guinea, and possibly the Philippines. Modern Nymphinae is only found in Australia. This may result from a requirement of frost-free climates, which were more widespread in the past.

Fossil dragonflies (Odonata: Anisoptera) from the early Eocene Okanagan Highlands, western North America

We describe the first dragonflies (Odonata: Anisoptera) from the early Eocene Okanagan Highlands of far-western North America from nine fossils. Six are assigned to five species in four new, named genera of Aeshnidae: Antiquiala snyderaenew genus and species, Idemlinea versatilisnew genus and species, Ypshna brownleeinew genus and species, Ypshna latipennatanew genus and species, and Eoshna thompsonensisnew genus and species; we treat one as Aeshnidae genus A, species A; one is assigned to Gomphidae: Auroradraco eosnew genus and species; and we treat a ninth, fragmentary fossil of unknown family affinity as Anisoptera indeterminate genus A, species A, which represents a seventh genus and eighth species. The dominance of Aeshnidae is consistent with other Paleocene and Eocene fossil localities. Auroradraco eos is the only fossil Gomphidae in the roughly 66-million-year gap between occurrences in mid-Cretaceous Burmese amber and the early Oligocene of France. Ypshna appears close to Parabaissaeshna ejerslevense from the early Eocene Fur Formation of Denmark; this is not surprising given Holarctic intercontinental connections at this time and a growing list of insect taxa shared between the Okanagan Highlands and the Fur Formation.

Early Eocene (Ypresian) birds from the Okanagan Highlands, British Columbia (Canada) and Washington State (USA)

We survey the known avian fossils from Ypresian (early Eocene) fossil sites of the North American Okanagan Highlands, mainly in British Columbia (Canada). All specimens represent taxa that were previously unknown from the Eocene of far-western North America. Wings from the McAbee site are tentatively referred to the Gaviiformes and would constitute the earliest fossil record of this group of birds. A postcranial skeleton from Driftwood Canyon is tentatively assigned to the Songziidae, a taxon originally established for fossils from the Ypresian of China. Two skeletons from Driftwood Canyon and the McAbee site are tentatively referred to Coliiformes and Zygodactylidae, respectively, whereas three further fossils from McAbee, Blakeburn, and Republic (Washington, USA) are too poorly preserved for even a tentative assignment. The specimens from the Okanagan Highlands inhabited relatively high paleoaltitudes with microthermal climates (except Quilchena: lower mesothermal) and mild winters, whereas most other Ypresian fossil birds are from much warmer lowland paleoenvironments with upper mesothermal to megathermal climates. The putative occurrence of a gaviiform bird is particularly noteworthy because diving birds are unknown from other lacustrine Ypresian fossil sites of the Northern Hemisphere. The bones of the putative zygodactylid show a sulphurous colouration, and we hypothesize that this highly unusual preservation may be due to the metabolic activity of sulphide-oxidizing bacteria.

The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America

One new genus of Inocelliidae (Raphidioptera) with one new species and one undetermined specimen is described from the Eocene of North America: Paraksenocellia borealis new genus, new species from the early Eocene (Ypresian) Okanagan Highlands shale at Driftwood Canyon, British Columbia, Canada (a forewing), and Paraksenocellia species from the middle Eocene (Lutetian) of the Coal Creek Member of the Kishenehn Formation, northwestern Montana, United States of America (a hind wing). These are the oldest records of the family. The new genus possesses many character states that are rare in Inocelliidae, e.g., a very long pterostigma extending to ScP in both the forewings and hind wings; the forewing subcostal space has three crossveins; the forewing and hind wing AA1 are deeply forked; the crossvein between CuA and CuP is located far distad the crossvein 1r-m. Paraksenocellia is confidently a member of the Inocelliidae, as it possesses a proximal shift of the basal crossvein 1r-m (connecting R and M) in the forewing and the loss of the basal crossvein 1r-m in the hind wing, both apomorphies of the family. It shares some character states with the Mesozoic Mesoraphidiidae, which we consider to be mostly stem-group plesiomorphies.