Stolleagrion foghnielseni (Odonata, Cephalozygoptera, Dysagrionidae) gen. et sp. nov.: a new odonatan from the PETM recovery phase of the earliest Ypresian Fur Formation, Denmark.

We describe the new genus and species Stolleagrion foghnielseni n. gen. et sp. from the Fur Formation in northwestern Denmark based on a single fossil wing. This is the first odonatan described from the earliest part of the PETM recovery phase of the early Eocene. A combination of nine wing character states are considered to be diagnostic of the Dysagrionidae Cockrell only together with the cephalozygopteran head; however, the combination of these nine plus the presence of Ax0 is also diagnostic without the head. By this, we assign Stolleagrion foghnielseni to the Dysagrionidae and reassess the position of other odonates previously treated as cf. Dysagrionidae.

The damselfly genus Furagrion Petrulevicius et al. (Odonata, Zygoptera) from the early Eocene Fur Formation of Denmark and the dysagrionoid grade.

The earliest Eocene odonate genus Furagrion Petrulevicius et al. from the Danish Fur Formation is revised based on eighteen specimens, two of which apparently have been lost since their publication. The holotype of Phenacolestes jutlandicus Henriksen, type species of Furagrion, is incomplete and lacks the characters currently used to differentiate species, genera and higher taxa in Odonata. We, therefore, propose that the holotype is set aside and a recently discovered nearly complete Fur Formation fossil is designated as neotype. Furagrion possesses all of the nine wing character states currently used along with head shape for diagnosing the Dysagrionidae; however, Furagrion has a characteristically zygopteran head, not the distinctive head shape of the suborder Cephalozygoptera. We, therefore, treat it as a zygopteran unassigned to family. These nine wing character states appear in different combinations not only in various Zygoptera and Cephalozygoptera, but also in the Frenguelliidae, an Eocene family of Argentina that may represent an unnamed suborder. We recognise these taxa as constituting a dysagrionoid grade, in which these character states appear either convergently or as symplesiomorphies. Furagrion morsi Zessin is synonymized with Phenacolestes jutlandicus Henriksen, syn. nov. and Morsagrion Zessin with Furagrion Petrulevicius, Wappler, Wedmann, Rust, and Nel, syn. nov.

Cimbicidae (Hymenoptera, 'Symphyta') in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands.

We erect the Cenocimbicinae, a new subfamily of Cimbicidae (Hymenoptera, Symphyta), restricted to the Selandian Menat Formation of France, the oldest occurrence of the family, and the Ypresian Okanagan Highlands of far-western North America. We describe new taxa from the Okanagan Highlands: Allenbycimbex morrisae gen. et sp. nov. and Leptostigma n. gen. with seven new species: L. alaemacula n. sp., L. brevilatum n. sp., L. fasciatum n. sp., L. longiclava n. sp., L. longipallidum n. sp., L. longitenebricum n. sp., and L. proxivena n. sp. We revise the Cimbicidae from the Ypresian Green River Formation and the Priabonian Florissant Formation, both in Colorado, USA. The oldest fossil of a modern cimbicid subfamily appears with a single pachylostictine specimen in the Green River Formation, and all Cimbicidae are in modern subfamilies after the Ypresian (we did not examine one larva known from Priabonian Baltic amber). Pseudocimbex clavatus Rohwer 1908 from the Florissant Formation is not a cimbicid; we treat it as Tenthredinoidea incertae sedis. We transfer Cimbex vetusculus Cockerell to Floricimbex n. gen.

A revision of Chrysopidae (Neuroptera) from the late Eocene Florissant Formation, Colorado, with description of new species.

The green lacewings (Neuroptera: Chrysopidae) of the late Eocene Florissant Formation (Colorado, USA) are revised. Thirteen species in six genera of Nothochrysinae are recognized, including four new species: Archaeochrysa cockerelli sp. nov., Cimbrochrysa americana sp. nov., C. major sp. nov., and Lithochrysa meyeri sp. nov. The validity of the genus Dyspetochrysa Adams, 1967 is tenuous; it might be a synonym of Archaeochrysa Adams, 1967. Lithochrysa Carpenter, 1935 and L. ferruginea (Cockerell, 1909) are considered a valid genus and species. A lectotype and paralectotype of Tribochrysa firmata Scudder, 1890 are designated. The genus Cimbrochrysa Schlter, 1982 is recorded from North America for the first time. Eocene chrysopid assemblages are briefly analyzed. The late Eocene Florissant assemblage is less diverse than those of early Eocene North America, and greatly differs from late Eocene European assemblages.

Danowhetaksa gen. nov. with two species from the early Eocene lst Formation from Denmark, the first Palearctic Whetwhetaksidae (Odonata: Cephalozygoptera).

We propose Danowhetaksa n. gen. (Odonata: Whetwhetaksidae) with two new species: D. birgitteae n. gen. et sp. and D. rusti n. gen. et sp. from the earliest Ypresian Stolleklint clay of the lst Formation in northwestern Denmark. Whetwhetaksidae has previously been known only from the Ypresian Okanagan Highlands of far-western North America, the new records are, therefore, the first from the Palearctic Region.

Kishenehna prima, a new genus and species of darner dragonfly (Odonata, Aeshnidae, Gomphaeschninae) from the early middle Eocene Kishenehn Formation of Montana, USA.

We describe the darner dragonfly Kishenehna prima n. gen. and sp. (Odonata, Aeshnidae, Gomphaeschninae) based on a well-preserved, nearly complete female hind wing from the Lutetian Coal Creek Member of the Kishenehn Formation, northwestern Montana, USA. Kishenehna is morphologically close to the late Paleocene genus Alloaeschna Wighton Wilson of Alberta, Canada. This is the first dragonfly (Anisoptera) described from the Kishenehn Formation and the first from the Lutetian of the Western Hemisphere.

The Palaeorehniidae (Orthoptera, Ensifera, Zeuneropterinae), and new taxa from the early Eocene Okanagan Highlands, western North America.

We revise the subfamily Zeuneropterinae (Orthoptera, Ensifera) and treat it as the family Palaeorehniidae Zeuner stat. nov. We restore the late Eocene Palaeorehnia Cockerell from Florissant, Colorado, USA to the taxon and describe two new genera and species in it: Ypopteron nicola n. gen., n. sp. from Quilchena, British Columbia, Canada and Republicopteron douseae n. gen., n. sp. from Republic, Washington, USA. Given current knowledge, Palaeorehniidae appears intermediate between the Stenopelmatoidea and the Hagoidea (s.s.).

The second North American fossil horntail wood-wasp (Hymenoptera: Siricidae), from the early Eocene Green River Formation.

Eoteredon lacoi gen. et sp. nov., is described from the early Eocene Green River Formation in Wyoming, the second fossil siricid genus and species described from North America. We propose Eoteredon as sister to the genus Teredon, whose sole species is one of the rarest of extant Siricidae. The majority of siricids today inhabit temperate Northern Hemisphere forests; Teredon cubensis, however, is one of its few species that live in megathermal tropical lowlands. The Eocene forest that Eoteredon inhabited had a mesothermal to megathermal climate. We place Eoteredon in the context of broad Cenozoic climate change.

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.

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.

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).

The second genus and species of the extinct neuropteroid family Corydasialidae, from early Eocene McAbee, British Columbia, Canada: do they belong to Megaloptera?

Ypresioneura obscura gen. et sp. nov. from the early Eocene (Ypresian) McAbee locality (Canada, British Columbia) is described. It is assigned to the extinct neuropteroid family Corydasialidae, as the second known genus and species. The Corydasialidae was previously known only from late Eocene (Priabonian) Baltic amber. It was originally assigned to the Megaloptera, but the character states that support this conclusion are not strongly diagnostic. There is still doubt as to whether this family belongs to Megaloptera or to the Neuroptera. If this is a megalopteran (which we favor), it is the first specimen of the order from the insect-rich Okanagan Highlands series of Ypresian localities, which occur sporadically across the southern interior of British Columbia, Canada into north-central Washington, USA.

An unusual new fossil genus probably belonging to the Psychopsidae (Neuroptera) from the Eocene Okanagan Highlands, western North America.

The new genus and species Ainigmapsychops inexspectatus gen. et sp. nov. is described from the early Eocene Okanagan Highlands locality at Republic, Washington, U.S.A. We preliminarily assign it to the Psychopsidae; however, its venation is unusual within this family, particularly by its pectinate branches of AA1 originating at a steep angle, a character state more suggestive of the Osmylidae. 

A revision of the late Eocene snakeflies (Raphidioptera) of the Florissant Formation, Colorado, with special reference to the wing venation of the Raphidiomorpha.

The snakeflies (Raphidioptera) of the late Eocene Florissant Formation (Colorado, USA) are revised. Seven species of Raphidiidae are assigned to three genera, i.e., Megaraphidia Cockerell, 1907, Archiraphidia Handlirsch, 1910, and Florissantoraphidia gen. nov. Dictyoraphidia Handlirsch, 1910 is assigned to Baissopteridae, a first Cenozoic record of the family. Archiraphidia tumulata (Scudder, 1890), A. tranquilla (Scudder, 1890) and A.? somnolenta (Scudder, 1890), stat. res. are treated as distinct species, and A. eventa (Scudder, 1890) as a new synonym of A. tranquilla. The lectotype of A. eventa is designated. 'Raphidia' exhumata Cockerell, 1909 is transferred to Megaraphidia. 'Raphidia' mortua Rohwer, 1909 and 'R.' funerata Engel, 2003 constitute Florissantoraphidia gen. nov. Our findings support the treatment of the single Florissant species of Inocelliidae as preliminary assigned to Fibla Navás, 1915. We examine venational synapomorphies of Raphidiomorpha and provide a new diagnosis for it based on these characters. We evaluate putative derived venational character states of 'Neoraphidioptera' (Inocelliidae and Raphidiidae), finding no clear synapomorphy supporting its validity; these families may nest separately within a paraphyletic Mesoraphidiidae (s.l.). We provide diagnoses for the families occurring at Florissant (Baissopteridae, Inocelliidae and Raphidiidae) based on wing venation.

Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum.

Eocene climate and associated biotic patterns provide an analog system to understand their modern interactions. The relationship between mean annual temperatures and winter temperatures-temperature seasonality-may be an important factor in this dynamic. Fossils of frost-intolerant palms imply low Eocene temperature seasonality into high latitudes, constraining average winter temperatures there to >8 °C. However, their presence in a paleocommunity may be obscured by taphonomic and identification factors for macrofossils and pollen. We circumvented these problems by establishing the presence of obligate palm-feeding beetles (Chrysomelidae: Pachymerina) at three localities (a fourth, tentatively) in microthermal to lower mesothermal Early Eocene upland communities in Washington and British Columbia. This provides support for warmer winter Eocene climates extending northward into cooler Canadian uplands.

New species of Nymphites Haase (Neuroptera: Nymphidae) from the Middle Jurassic of China, with a redescription of the type species of the genus.

Nymphites priscus (Weyenbergh, 1869) from the Late Jurassic of Solnhofen (Germany), type species of the genus Nymphites Haase, 1890, is redescribed. The genus is assigned to the Nymphidae. The taxon Nymphitidae is not valid; it is an artificial aggregation of fossil genera. Two species new to science, one named, of Nymphites from the Middle Jurassic locality of Daohugou (Inner Mongolia, China) are described.

Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals.

Early Eocene land bridges allowed numerous plant and animal species to cross between Europe and North America via the Arctic. While many species suited to prevailing cool Arctic climates would have been able to cross throughout much of this period, others would have found dispersal opportunities only during limited intervals when their requirements for higher temperatures were met. Here, we present Titanomyrma lubei gen. et sp. nov. from Wyoming, USA, a new giant (greater than 5 cm long) formiciine ant from the early Eocene (approx. 49.5 Ma) Green River Formation. We show that the extinct ant subfamily Formiciinae is only known from localities with an estimated mean annual temperature of about 20°C or greater, consistent with the tropical ranges of almost all of the largest living ant species. This is, to our knowledge, the first known formiciine of gigantic size in the Western Hemisphere and the first reported cross-Arctic dispersal by a thermophilic insect group. This implies intercontinental migration during one or more brief high-temperature episodes (hyperthermals) sometime between the latest Palaeocene establishment of intercontinental land connections and the presence of giant formiciines in Europe and North America by the early middle Eocene.