Paleofauna of the Eocene Okanagan Highlands

Not to be confused with Paleoflora of the Eocene Okanagan Highlands.

The paleofauna of the Eocene Okanagan Highlands consists of Early Eocene arthropods, vertebrates, plus rare nematodes and molluscs found in geological formations of the northwestern North American Eocene Okanagan Highlands. The highlands lake bed series' as a whole are considered one of the great Canadian Lagerstätten. The paleofauna represents that of a late Ypresian upland temperate ecosystem immediately after the Paleocene-Eocene thermal maximum, and before the increased cooling of the middle and late Eocene to Oligocene. The fossiliferous deposits of the region were noted as early as 1873, with small amounts of systematic work happening in the 1880-90s on British Columbian sites, and 1920-30s for Washington sites. Focus and more detailed descriptive work on the Okanagan Highlands site started in the last 1970's. Most of the highlands sites are preserved as compression-impression fossils in "shales", but also includes a rare permineralized biota and an amber biota.

Driftwood

Horsefly

McAbee

Hat Creek

Falkland

Quilchena

Princeton

Republic

class=notpageimage|

Major Eocene Okanagan Highlands sites

## Extent[edit]

The 1,000 km (620 mi) series of lacustrine deposits are located across the Central British Columbia, Canada southeast to northern central Washington state, United States. grouped informally into "Northern", "Central", and "Southern" sites.[1] The Northern sites consist of unnamed Ootsa Group formations which outcrop as the "Driftwood shales" near Smithers, British Columbia, the "Horsefly shales", of an unnamed formation and unnamed group which outcrop around Horsefly, British Columbia,[1] and possibly sites now considered lost in the Quesnel, British Columbia area,[2] The Central sites represent Kamloops Group formations with the McAbee Fossil Beds, Tranquille River site and Falkland site, all in the Tranquille Formation, the Quichena site and Stump Lake site in the Coldwater Beds and outcrops of the Chu Chua Formation near Barriere, British Columbia. The Southern sites include the Princeton Group Allenby Formation sites surrounding Princeton, British Columbia, such as "Nine Mile Creek", "One Mile Creek", "Pleasant Valley", "Thomas Ranch", "Vermilian Bluffs", and "Whipsaw Creek". The Penticton Groups Kettle River, Marama and Marron Formations in the Boundary District along the Canada-United States border are closely correlated with the Klondike Mountain Formation across the border.[3] The most southerly of the Okanagan Highlands lakes, the Klondike Mountain Formation in Northern Ferry County, Washington include the "Boot Hill site", "Corner Lot site", "Gold Mountain site", "Knob Hill site", and "Mount Elizabeth site".[1]

There is debate as to the affiliation of the, potentially lost, Quesnel sites with the Greater Okanagan Highlands. Archibald et al. (2018) in a monograph of the Highlands Hymenoptera families included them as part of the series.[4] However the certainty for the placement was questioned earlier by Eberle et al. (2017)[5] and Archibald and Cannings (2022) who opted to tentatively exclude Quesnel from the highlands while discussing the history of field colleting in the region.[2]

## Paleofauna[edit]

C. bittaciformis

C. brooksi

Cimbrophlebia species
Fur Formation (left), Okanagan Highlands (right)

The Okanagan highlands represent a snapshot of lake, wetlands, and montane forest animal life which existed approximately 15 million years ago after the Cretaceous–Paleogene extinction event. The temperate upland lakes hosted insects, fish, birds, and mammals with the notably well preserved megafossils often retaining insect colour patterns, gnat wing membrane hairs, and whole bird feathers.[6] In some cases the fine detail preservation of soft parts allows for the preservation of internal anatomy.[7]

Archibald and Makarkin (2006) suggested the disjunct distribution of genera between the Danish western Limfjord coasts Fur Formation and the Okanagan highlands may have been enabled by rising crust elevations in the northern Atlantic region and subsequent increase in landmass during the Late Paleocene which linked Northern Europe with Greenland until at least the Early Eocene.[8] Several land bridge routes may have acted as migration corridors for biotic interchange, the northern De Geer land bridge from Fennoscandia to North America via northern Greenland, and the southern Thulean land bridge from northern Britain though the Faroe Islands and then Greenland and North America. Several insect genera share disjunct distributions between the highlands and Limfjord including the mecopteran Cimbrophlebia, the giant lacewing Palaeopsychops, the green lacewing Protochrysa, the bull dog ant Ypresiomyrma.[8]

The Hat Creek Amber deposits in the central region provide evidence for small and microbiotic elements of the Okanagan Highlands forests though entombed organisms such at terrestrial nematodes and microwasps that otherwise would likely not be preserved in the lake environments.[9] The highlands as a whole have been described as one of the "Great Canadian Lagerstätten"[6] based on the diversity, quality and unique nature of the 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.[6] David Grimaldi et al. (2018) during discussion of inclusions in Alaskan Chickaloon amber, noted the Okanagan Highlands record of latitudinal extinctions, specifically the modern southern hemisphere endemic groups Eomeropidae mecopterans and Myrmeciinae bulldog ants.[10]

## Lithology[edit]

The majority of the lake deposits are compression fossils in lake bed sediments noted for both the paleofauna and paleofloras, with an additional pair of important non-compression biotas. A permineralized chert flora, the Princeton Chert is found along the Similkameen River interbedded with coal deposits of the Ashnola shale unit, Allenby Formation known for anatomically preserved plants.[11] In the Central sites, subbituminous coal of the Hat Creek Coalfield around Hat Creek hosts an entombment biota, the Hat Creek amber, which preserves highlands faunal elements that are not found in the compression biotas.[9][4] Initial discussion of the amber presented by George Poinar, Jr. et al. (1999) suggested the Hat Creek amber producing tree was likely to be an araucarian tree in the genus Agathis, based on unreported magnetic resonance spectroscopic analysis and earlier reports of the genus in Mesozoic Canada.[12] A purported occurrence of Araucaria at the McAbee site was used as additional support for the ambers origin.[13] The Agathis origin for Canadian Mesozoic amber was later called into question by Ryan McKellar and Alexander Wolfe (2010) based on a lack of any araucarian macrofossil history in the northern hemisphere[12] the McAbee fossils having been already reidentifed as from the cupressaceous Cunninghamia.[13] Based on Fourier-transform infrared spectroscopic analysis and associated amber inclusion fossils, they suggested the Mesozoic ambers of Canada to be from the extinct cupressaceous genus Parataxodium.[12] The origin of the Hat Creek ambers was further noted as likely from a cupressaceous source by Grimaldi et al. (2018) who call out a primary floral component of the host coal being Metasequoia and that the coeval Puget Group Tiger Mountain amber of Washington state is also of Metasequoia origins. They hypothesize that the major amber producing plant of the Paleocene Pacific Northwest forests as Metasequoia, but note that further investigation of Chickaloon, Hat Creek, Coalmont, and Tiger Mountain ambers would be needed.[10]

## Mollusks[edit]

Mollusks are a rare component of the highlands, usually being mentioned only in passing, such as by Mark Wilson (1977, 1978),[14][15] and with fossils being reported from three sites only. A series of species were described from several Allenby Formation sites around Princeton by Russell (1957), who documented 4 gastropod species, and tentatively identified to genus another gastropod and a bivalve.[16] Additional unidentified small bivalve fossils were mentioned from the Pleasant Valley site by Wilson (1977) and the Quilchena site by Wilson (1987),[17] while unidentified gastropods were briefly mentioned by Kathleen Pigg et al. (2018).[18]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Hydrobiidae

†Micropyrgus

†Micropyrgus camselli[16]

* Princeton

A hydrobiid mud snail

Lymnaeidae

Stagnicola

†Stagnicola tulameenensis[16]

* Princeton

A lymnaeine pond snail

Physidae |

Aplexa

†Aplexa ricei[16]

* Princeton

An aplexine bladder snail

Physa

†Physa saxarubrensis[16]

* Princeton

A physine bladder snail

Planorbidae |

Ferrissia

†Ferrissia arionoides[16]

* Princeton

An ancylinine ramshorn snail

Gyraulus?

Indeterminate[16]

* Princeton

A possible planorbinine ramshorn snail
Not described to species

Sphaeriidae

Sphaerium?

Indeterminate[16]

* Princeton

A possible sphaeriine fingernail clam
Not described to species

Unidentified

Unidentified

Unidentified[14][17]

* Quilchena
* Princeton

Unidentified freshwater bivalves.

Unidentified

Unidentified

Unidentified[18]

* Republic

Unidentified freshwater gastropods.

|  Undescribed gastropod

## Nematodes[edit]

In the initial description of Hat Creek Amber, Poinar et al. make note of nematode specimens found in the deposit, with a brief commentary regarding them as the oldest terrestrial free-living nematode fossils to have been found up to that point, but did not give any specific taxonomic identification beyond that.[9]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Unidentified

Unidentified

Unidentified[9]

* Hat Creek

Unidentified terrestrial specimens in amber

## Arachnids[edit]

A single arachnid has been described fully from the Okanagan highlands, the Nursery web spider Palaeoperenethis thaleri, known from an adult male. This spider was likely aquatic as are the other members of the family, and based on morphological similarities, it was possibly closer in relation to African and Asian species in the Perenethis genus group then to the only modern genus in British Columbia Dolomedes.[19] Another spider specimen, University of Alberta 5007 was noted by Wilson (1977) from the Kamloops area, while various spiders have been attributed to the McAbee fossil beds.[20] Undescribed male and female orb-web spiders were figured by Wehr (1998) from the Princeton area.[21] Unspecified spider compression fossils were mentioned as occurring in passing by David Greenwood et al. (2005) while discussing the increasing taxonomic richness of the highlands, but without specific site information.[22] Additionally, undiscussed amber fossils were mentioned by Poinar et al. (1999) in their initial report of Hat Creek amber inclusions.[9] Other arachnid evidence has been recovered in the form of fossil hymenopterans placed in families known predate or parasitize spiders. A diverse undescribed fauna of the "parasitoid" wasp family Ichneumonidae is known, some species of which are known to parasitize eggs or adult spiders. Another family, Sphecidae, which is a documented opportunistic predator of spiders and certain insets is known from a few isolated fossils at McAbee and Republic. Lastly the vespoid family Pompilidae has been found at both McAbee and Republic. This family, known as spider wasps, are behaviorally specialized as predators of spiders and a few other arachnids, provisioning newly laid eggs with a single spider as a larder to feed on while developing.[4] Galling preserved on Acer species leaf fossils has been attributed to mites in the family Eriophyidae.[23]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Araneidae

Unidentified

Unidentified[21]

* Princeton

An orb-weaver spider
Not described to genus/species

Pisauridae

Palaeoperenethis

Palaeoperenethis thaleri[19]

* Horsefly

A Nursery web spider

Eriophyidae

Unidentified

Unidentified[23]

* Republic

Trace fossils
Eriophyid gall mite galling on Acer leaves
Not described to genus/species

Unidentified  | Unidentified  |

Unidentified[24][20]

* "Kamloops sites"
Tranquille or Falkland
* McAbee

Spider Compression fossils from Kamloops area locations
Not described to family/genus/species

Unidentified[24]

* McAbee

Possible Araneidae? or Tetragnathidae? fossils
Not described to family/genus/species

Unidentified[22]

* Unspecified

Spider compression fossils from unspecified highlands locations
Not described to family/genus/species

Unidentified[9]

* Hat Creek

Spider specimens in amber
Not described to family/genus/species

## Crustaceans[edit]

The earliest report of Crayfish from the highlands was by Wesley Wehr and Lisa Barksdale (1995). In a short Washington Geology article they reported the first identified occurrence of feathers from the Klondike Mountain Formation and crayfish from both there and the McAbee site. At that time, the moulted carapace section from Republic was not identified further than as a freshwater crayfish. The McAbee specimen was tentatively identified, from photograph, as being a possible Procambarus species fossil by malacologist Rodney M. Feldmann.[25] Subsequently an additional series of over ten fossils were recovered from McAbee and described in 2011 as Aenigmastacus crandalli by Feldmann, Carrie Schweitzer, and John Leahy. A. crandalli was placed in the southern hemisphere superfamily Parastacoidea based on several morphological characters, and they noted this species to be the only northern hemisphere member of the superfamily.[26]

At the Quilchena site, brief mention was reported in 2016 of ostracod fossils, though no further discussion or description has happened.[27]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Parastacidae

Aenigmastacus

Aenigmastacus crandalli[26]

* McAbee

A parastacid crayfish.
first illustrated as Procambarus sp. (1995)[25][26]

unidentified

Unidentified

unidentified[25]

* Republic

An unidentified crayfish

|  Crayfish moult
undescribed

unidentified

Unidentified

unidentified[28][27]

* Quilchena

Unidentified ostracod shells

## Insects[edit]

### Blattodea[edit]

In the initial description of Hat Creek Amber, Poinar et al. make note of a single adult Corydiinae cockroach specimen found in the amber, with a brief commentary on the modern tropical-subtropical distribution of that subfamily and a lack of any native cockroach species in western Canada, but did not give any specific taxonomic identification for the specimen beyond that.[9]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Blaberidae

Undescribed

Undescribed[27][22]

* Driftwood
* McAbee
* Quilchena
* Republic

Undescribed Diplopterine cockroaches.

Blattoidae

Undescribed

Undescribed[29]

* Republic

A blattoidean cockroach
Not described to genus/species

|  Undescribed Blattoidea

Corydiidae

Unidentified

Unidentified[9]

* Hat Creek

An undescribed corydiine cockroach

Hodotermitidae

Unidentified

Unidentified[20]

* McAbee

Harvester termites
Not described to genus/species

Mastotermitidae

Undescribed

Undescribed[24][14][22][20]

* Driftwood
* Falkland
* McAbee
* Horsefly
* Princeton

Darwin termites
Not described to genus/species

undescribed

Undescribed

Undescribed[29]

* Republic

Undescribed termites of uncertain affiliation

|  undescribed isopteran

### Coleopterans[edit]

The order Coleoptera is divided into four major lineages, Adephaga, Archostemata, Myxophaga, and Polyphaga, with the last group being the most species diverse of the four. Hat Creek amber has provided one fully described beetle species Prionocerites tattriei,[30] which is known from a larval stage specimen first reported by Poinar et al. (1999).[9] The species and genus were the first North American taxon from the family to be described.[30]

#### Adephaga[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Carabidae |

Amara

†Amara paleomelas[31]

* Quilchena

A sun beetle
First described as Nebria paleomelas[32]

|  Amara paleomelas
(1890 illustration)

Cf. Amara

Undescribed[27]

* Quilchena

A sun beetle relative

Undescribed

Undescribed[29][33][27][20]

* McAbee
* Quilchena
* Republic

Undescribed ground beetles.

|  Unidentified Carabidae

Dytiscidae

Undescribed

Undescribed[29]

* Republic

A diving beetle

Undescribed

Undescribed

Undescribed[33]

* Princeton

A caraboid superfamily beetle
Displays traits similar to both tiger beetles and scarabs
not identified to genus or species

#### Archostemata[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Cupedidae

Undescribed

Undescribed[20]

* McAbee

Reticulated beetles
Not described to genus/species

#### Polyphaga[edit]

##### Cucujiformia[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Brentidae |

†Eoceneithycerus

†Eoceneithycerus carpenteri[34]

* Republic

An Ithycerinae weevil

|  Eoceneithycerus carpenteri

†Ithyceroides

†Ithyceroides klondikensis[35]

* Republic

An Ithycerinae weevil

Cerambycidae

Undescribed

Undescribed[29][33][20]

* McAbee
* Quilchena?
* Republic

long-horn beetles
Not described to genus/species

Chrysomelidae |

†Cryptocephalites

†Cryptocephalites punctatus[36]

* Princeton

A leaf beetle

|  Cryptocephalites punctatus
(1895 illustration)

Galerucella

†Galerucella picea[32]

* Princeton

A leaf beetle

|  Galerucella picea
(1890 illustration)

Caryobruchus–Speciomerus genus group

Undescribed[37]

* Driftwood?
* McAbee
* Quilchena
* Republic

palm beetles, originally identified as cf. tribe Caryopemina.[27]

Undescribed

Undescribed[29][20]

* McAbee
* Republic

leaf beetles
Not described to genus/species

Curculionidae

Undescribed

Undescribed[29][33][27][20]

* Horsefly
* McAbee
* Quilchena
* Republic

Undescribed weevils

Mordellidae

Undescribed

Undescribed[20]

* McAbee

Pintail beetles
Not described to genus/species

Prionoceridae

†Prionocerites

†Prionocerites tattriei[30]

* Hat Creek

A Prionocerid beetle
Desccribed from a larval specimen
First reported by Poinar et al. (1999 Fig.7) as "A beetle larva"

Tenebrionidae

Tenebrio

†Tenebrio primigenius[32]

* Princeton

A darkling beetle

|  Tenebrio primigenius
(1890 illustration)

Cf. Cleridae

Undescribed

Undescribed[20]

* McAbee

Undescribed checkered beetle relatives

Cf. Erotylidae

Undescribed

Undescribed[27]

* Quilchena

Undescribed pleasing fungus beetle relatives.

Unidentified

Undescribed

Undescribed[33]

* Republic

A cucujiform beetle
not identified to family

##### Elateriformia[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Buprestidae | Buprestis |

†Buprestis saxigena[32][36]

* Quilchena

A jewel beetle

|  Buprestis saxigena
(1890 illustration)

†Buprestis sepulta[32]

* Quilchena

A jewel beetle

|  Buprestis sepulta
(1890 illustration)

†Buprestis tertiaria[32]

* Quilchena

A jewel beetle

|  Buprestis tertiaria
(1890 illustration)

Cantharidae

Undescribed

Undescribed[33]

* McAbee(?)
* Princeton

A soldier beetle
not identified to genus or species

Elateridae |

Ligmargus

†Ligmargus terrestris[38]

* Quilchena

A click beetle
First described as Cryptohypnus? terrestris[32]

|  Ligmargus terrestris
(1890 illustration)

Limonius

†Limonius impunctus[36][38]

* Princeton

a click beetle

|  Limonius impunctus
(1895 illustration)

†Elaterites

undescribed[36]

* Quilchena

A click beetle.

|  Elateridae sp. indet
(1890 illustration)

Undescribed

Undescribed[29][33][20]

* McAbee
* Princeton
* Republic

undescribed click beetles

##### Scarabaeiformia[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Lucanidae

Undescribed

Undescribed[29][33]

* Republic

A stag beetle

Passalidae

Undescribed

Undescribed[20]

* McAbee

Bess beetles
Not described to genus/species

Trogidae

Trox

†Trox oustaleti[32]

* Princeton

A hide beetle

|  Trox oustaleti
(1890 illustration)

Undescribed

Undescribed

Undescribed[33][27]

* Quilchena

Undescribed scarab superfamily beetles.

##### Staphyliniformia[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Hydrophilidae

Cercyon?

†Cercyon? terrigena[32]

* Quilchena

A water scavenger beetle

|  Cercyon? terrigena
(1890 illustration)

Staphylinidae

Undescribed

Undescribed[27]

* Quilchena

Undescribed Omaliinae rove beetles.

Unidentified

Undescribed

Undescribed[33]

* Republic

A possible staphylinoid beetle
not described

### Dermapterans[edit]

Earwig fossils were first noted from republic by paleoentomologist Standley Lewis (1992) in his initial report of the insect diversity at Republic. He noted the fossils to be some of the oldest Eocene demapterans in North America at that time and figured one undescribed specimen consisting of a females abdomen section and cerci.[29] Lewis (1994) tentatively identified the earwigs as members of family Forficulidae based on the shape of the cerci, and illustrated four female fossils, identified as such from the simple straight nature of the cercus. Lewis also suggested two different species were present, based on the differing lengths of the female cerci.[39]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

?Forficulidae | Undescribed  |

"Forficulid species 1"[39]

* Republic

A forficulid? earwig species with long cerci

|  "Forficulid species 1"
undescribed

"Forficulid species 2"[39]

* Republic

A forficulid? earwig species with short cerci

|  "Forficulidae species 2"
undescribed

Unidentified[20]

* McAbee

A possible forficulid earwig
Not described to genus/species

Forficulidae

Undescribed

Undescribed[27]

* Quilchena

A forficuline earwig of undetermined placement

### Dipterans[edit]

The most common animal fossils at many of the highlands sites are bibionid march flies,[22] with over twenty species from the genera Penthetria and Plecia described.[40] The modern diversity of the family is greatest in lower latitudes, and Plecia only reaches northward to the warm temperate areas of southeastern North America.[22] In the initial description of Hat Creek Amber, Poinar et al. make note of dipteran inclusions found in the deposit but did not give any specific taxonomic identification of taxa or illustrate any specimen.[9]

#### Brachycera[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Agromyzidae

Undescribed

Undescribed[41]

* Princeton

Trace fossils
Agromyzid leaf mining
Not described to genus/species

Dolichopodidae

Microphor

†Microphor defunctus[42]

* Princeton

A long-legged fly
First described as Microphorus defunctus (1910),[43]
Moved to Microphor defunctus (1994)[42]

|  Microphor defunctus
(1910 illustration)

Empididae

Undescribed

Undescribed[24][33]

* Driftwood
* Horsefly
* Republic

dagger flies
Not described to genus/species

Pipunculidae |

†Metanephrocerus

†Metanephrocerus belgardeae[44]

* Republic

A big headed fly

|  Metanephrocerus belgardeae

Indeterminate

indeterminate[27][44]

* Quilchena

Big headed flies,
too incomplete to describe to genus

Rhagionidae

Undescribed

Undescribed[33]

* Quilchena

snipe flies
Not described to genus/species

Syrphidae

Unidentified

Unidentified[24][27][20][45]

* Driftwood
* Horsefly
* McAbee
* Quilchena
* Republic

hover flies or, less likely, thick headed flies
Not described to genus/species

#### Nematocera[edit]

The highlands have been noted for the number of Bibionid taxa named in the early work on British Columbian sites. Over 25 unique species have been ascribed to the fossils, with the current count sitting at 22, but doubt has been raised as to the true number of species present and around the generic affinities. The first species was described by Scudder (1879 from the Allenby Formation, while the largest number of species were named by Handlirsch (1910). Following the practice of the time, both Scudder and Handlirsch placed their species in the genus Penthetria. Harrington Molesworth Anthony Rice (1959) reviewed the British Columbian bibionids, notably deeming the majority of species as belonging to Plecia or an undescribed extinct genus and not Penthetria.[40] This placement decision has been questioned however, with Giuseppe Gentilini (1991) asserting the majority of highlands species should be returned to Penthetria.[46] Rice, noted to be a "splitter",[14] also noted the large overlap between the morphology of two species groups and mused that larger collection samples may reveal each group to be single species. He called out in the species discussions the similarities between Plecia avus, P. canadensis, P. dilatata, P. pictipennis, P. pulchra, and P. transitoria, and the similarities between P. curtula, P. nana, P. pulla, and P. reducta.[40]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Bibionidae |

Penthetria?

†Penthetria? fryi[40]

* Driftwood
* Princeton

A possible penthetrian march fly

|  Penthetria (?) fryi
Penthetria |

†Penthetria intermedia[40]

* Driftwood

A plecian marchfly
First described as Mycetophaetus intermedius (1892),[47]
moved to Plecia intermedia (1959)[40] moved to Penthetria intermedia (1999)[48]

†Penthetria whipsawensis[40]

* Princeton

A penthetrian march fly

|  Penthetria whipsawensis
Plecia |

†Plecia angustipennis[40]

* Horsefly
* Quilchena

A plecian marchfly
First described as Penthetria angustipennis (1910),[43] moved to Plecia angustipennis (1959)[40]

|  Plecia angustipennis

†Plecia avus[40]

* Driftwood
* Tranquille
* Princeton

A plecian marchfly
First described as Penthetria avus (1910),[43] moved to Plecia avus (1959)[40]
Possibly synonymous with Pl. canadensis, Pl. dilatata, Pl. pictipennis, Pl. pulchra, and Pl. transitoria[40]

|  Plecia avus

†Plecia cairnesi[40]

* Driftwood
* Falkland

A plecian marchfly

|  Plecia cairnesi

†Plecia canadensis[40]

* Driftwood
* Quilchena?
* Princeton
* Tranquille

A plecian marchfly
First described as Penthetria canadensis (1910),[43] moved to Plecia canadensis (1959)[40]
Possibly synonymous with Pl. avus Pl. dilatata, Pl. pictipennis, Pl. pulchra, and Pl. transitoria[40]

|  Plecia canadensis

†Plecia curtula[40]

* Driftwood
* Horsefly
* Princeton

A plecian marchfly
First described as Penthetria curtula (1910),[43] moved to Plecia curtula (1959)[40]
Pe. avunculus (1910) considered a jr synonym (1959)
Possibly synonymous with Pl. nana, Pl. pulla, and Pl. reducta[40]

|  Plecia curtula

†Plecia dilatata[40]

* Horsefly
* Princeton

A plecian marchfly
First described as Penthetria dilatata (1910),[43] moved to Plecia dilatata (1959)[40]
Possibly synonymous with Pl. avus, Pl. canadensis, Pl. pictipennis, Pl. pulchra, and Pl. transitoria[40]

|  Plecia dilatata

†Plecia elatior[40]

* Princeton

A plecian marchfly
First described as Penthetria elatior (1910),[43] moved to Plecia elatior (1959)[40]

|  Plecia elatior

†Plecia kelownaensis[40]

* Driftwood
* Mission Creek

A plecian marchfly

|  Plecia kelownaensis

†Plecia minutula[40]

* Princeton

A plecian marchfly

|  Plecia minutula

†Plecia nana[40]

* Princeton

A plecian marchfly
First described as Penthetria nana (1910),[43] moved to Plecia nana (1959)[40]
Possibly synonymous with Pl. curtula, Pl. pulla, and Pl. reducta[40]

|  Plecia nana

†Plecia pictipennis[40]

* Princeton
* Quilchena

A plecian marchfly
First described as Penthetria pictipennis (1910),[43] moved to Plecia pictipennis (1959)[40]
Pe. lambei (1910), Pe. ovalis (1910), & Pe. separanda (1910) considered jr synonyms (1959)[40]
Possibly synonymous with Pl. avus, Pl. canadensis, Pl. dilatata, Pl. pulchra, and Pl. transitoria[40]

|  Plecia pictipennis

†Plecia platyptera[40]

* Horsefly

A plecian marchfly
First described as Penthetria platyptera (1910),[43] moved to Plecia platyptera (1959)[40]

|  Plecia platyptera

†Plecia pulchra[40]

* Princeton

A plecian marchfly
First described as Penthetria pulchra (1910),[43] moved to Plecia pulchra (1959)[40]
Possibly synonymous with Pl. avus, Pl. canadensis, Pl. dilatata, Pl. pictipennis, and Pl. transitoria[40]

|  Plecia pulchra

†Plecia pulla[40]

* Princeton

A plecian marchfly
First described as Penthetria pulla (1910),[43] moved to Plecia pulla (1959)[40]
Pe. brevipes (1910) considered a jr synonym (1959)
Possibly synonymous with Pl. curtula, Pl. nana, and Pl. reducta[40]

|  Plecia pulla

†Plecia reducta[40]

* Driftwood
* Horsefly
* Princeton

A plecian marchfly
First described as Penthetria reducta (1910),[43] moved to Plecia reducta (1959)[40]
Possibly synonymous with Pl. curtula, Pl. nana, and Pl. pulla[40]

|  Plecia reducta

†Plecia similkameena[32]

* Princeton
* Tranquille

A plecian marchfly
First described as Penthetria similkameena (1879)[32] moved to Plecia similkameena (1885)[49]

|  Plecia similkameena

†Plecia transitoria[40]

* Driftwood
* Horsefly
* Princeton
* Tranquille

A plecian marchfly
First described as Penthetria transitoria (1910),[43] moved to Plecia transitoria (1959)[40]
Pe. falcatula (1910) & Pe. fragmentum (1910) considered jr synonyms (1959)[40]
Possibly synonymous with Pl. avus, Pl. canadensis, Pl. dilatata, Pl. pictipennis, and Pl. pulchra[40]

|  Plecia transitoria

†Plecia tulameenensis[40]

* Princeton
* Tranquille

A plecian marchfly

|  Plecia tulameenensis

Undescribed

Undescribed[29]

* Republic

A march fly
Not described to genus/species

|  Bibionidae
undescribed

Bolitophilidae

Undescribed

Undescribed[45]

* Horsefly

A bolitophilid fungus gnat
Not described to genus/species

Cecidomyiidae

Undescribed

Undescribed[41]

* Princeton
* Republic

Trace fossils
Cecidomyiid midge galling on various host leaves
Not described to genus/species

|  Cecidomyiidae gall on Prunus

Cylindrotomidae

Unidentified

Unidentified[20]

* McAbee

long-bodied craneflies
Not described to genus/species

Keroplatidae

Undescribed

Undescribed[45]

* Horsefly

A possible keroplatid fungus gnat
Not described to genus/species

Limoniidae

Unidentified

Unidentified[20]

* McAbee

Limoniid craneflies
Not described to genus/species

Mycetophilidae |

Undescribed

Undescribed[45]

* Horsefly

A gnoristine fungus gnat
Not described to genus/species

Undescribed

Undescribed[45]

* Horsefly

A leiine fungus gnat
Not described to genus/species

Undescribed

Undescribed[45]

* Horsefly

A mycetophiline fungus gnat
Not described to genus/species

Undescribed

Undescribed[45]

* Horsefly

A mycomyiine fungus gnat
Not described to genus/species

Undescribed

Undescribed[45]

* Horsefly

A sciophiline fungus gnat
Not described to genus/species

Undescribed

Undescribed[24][29][33][27][20]

* Driftwood
* McAbee
* Quilchena
* Princeton
* Republic

fungus gnats unplaced to subfamily
Not described to genus/species

Ptychopteridae

†Etoptychoptera

†Etoptychoptera tertiaria[50]

* Princeton

A phantom cranefly

|  Etoptychoptera tertiaria
(1910 illustration)

Sciaridae

Unidentified

Unidentified[24][33]

* Driftwood
* Horsefly
* Quilchena

dark-winged fungus gnats
Not described to genus/species

Tipulidae |

Tipula

†Tipula tulameena[43]

* Princeton

A cranefly

|  Tipula tulameena
(1910 illustration)

Undescribed

Undescribed[24][29][33][27][20][45]

* Driftwood
* Horsefly
* McAbee
* Princeton
* Quilchena
* Republic

crane flies
Not described to genus/species

|  Undescribed Tipulidae

Trichoceridae

Unidentified

Unidentified[20]

* McAbee

winter craneflies
Not described to genus/species

Unidentified

Unidentified

Unidentified[9]

* Hat Creek

Unidentified dipteran specimens in amber

### Ephemeropterans[edit]

Lewis (1992) listed one species of Heptageniidae and three specimens that he did not place to family from Republic.[29] The next year Lewis and Wehr (1993) gave a slightly more detailed description of the specimens again identifying one to Heptageniidae, possibly in the genera Heptagenia or Stenonema.[51] The specimens were later examined by Nina D. Sinitchenkova (1999) who described one as a squaregill mayfly and the oldest member of the genus Neoephemera, confirmed the Heptageniidae identification but that it was unidentifiable to genus. The last specimen she confirmed as an ephemeropteran, but unidentifiable below order level.[52]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Heptageniidae

Indeterminate

Indeterminate[52][20]

* McAbee
* Republic

A flat headed mayfly
not identifiable to genus

|  Heptageniidae incertae sedis

Neoephemeridae

Neoephemera

†Neoephemera antiqua[52]

* Republic

A squaregill mayfly

### Hemipterans[edit]

Greenwood et al. (2005) briefly discussed the prevalence of Aphid fossils at highlands sites where the taphonomic factors allowed for fine detail preservation such as in the Driftwood shales.[22] Poinar et al. (1999) made note of hemipteran specimens found in Hat Creek Amber but did not give any specific taxonomic identification or illustrate any specimens.[9]

#### Auchenorrhyncha[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Aphrophoridae | Aphrophora |

Aphrophora angusta[43]

* Princeton

An aphrophorine spittle bug hindwing species.

|  Aphrophora angusta
(1910 illustration)

"Indeterminate"[53]

* Princeton

An aphrophorine spittle bug
not described to species.

Undescribed[29]

* Republic

An aphrophorid spittlebug
Not described to species.

|  Aphrophora species

†Palaeoptysma

†Palaeoptysma venosa[53]

* Princeton

A spittle bug

|  Palaeoptysma venosa
(1895 illustration)

†Palaphrodes

"Indeterminate"[49]

* Princeton

An aphrophorine spittle bug
not described to species.

†Petrolystra

Undescribed[29]

* Republic

An aphrophorid spittlebug
Not described to species.

†Ptysmaphora

†Ptysmaphora fletcheri[53]

* Princeton

A spittle bug

|  Ptysmaphora fletcheri
(1895 illustration)
Cercopidae | Cercopis |

†Cercopis grandescens[53]

* Princeton

A froghopper

|  Cercopis grandescens
(1895 illustration)

†Cercopis selwyni[32]

* Princeton

A froghopper

|  Cercopis selwyni
(1890 illustration)

†Cercopites

†Cercopites torpescens[53]

* Princeton

A froghopper

|  Cercopites torpescens
(1895 illustration)

†Dawsonites

†Dawsonites veter[53]

* Princeton

A froghopper

|  Dawsonites veter
(1895 illustration)

†Palecphora

Undescribed[49][29]

* Princeton
* Republic

A froghopper
not described to species.

†Stenecphora

†Stenecphora punctulata[53]

* Princeton

A froghopper

|  Stenecphora punctulata
(1895 illustration)

†Stenolocris

†Stenolocris venosa[53]

* Princeton

A froghopper

|  Stenolocris venosa
(1895 illustration)

Undescribed

Undescribed[24][29][27][20]

* Driftwood
* McAbee
* Quilchena
* Republic

cercopid froghoppers
Not described to genus/species.

|  undescribed Cercopidae
Cicadellidae |

Coelidia

†Coelidia columbiana[32]

* Princeton

A leafhopper

|  Coelidia columbiana
(1890 illustration)

Undescribed

Undescribed[24][27][20]

* Driftwood
* McAbee
* Quilchena

leafhoppers
Not described to genus/species.

Cicadidae

Undescribed

Undescribed[20]

* McAbee

True cicadas
Not described to Family/genus/species

Cixiidae

Undescribed

Undescribed[27]

* Quilchena

A cixiid planthopper
Not described to genus/species.

Fulgoridae

Enchophora

"Indeterminate"[53]

* Princeton

An enchophorine lantern bug
not described to species.

Ricaniidae

Undescribed

Undescribed[27][20]

* McAbee
* Quilchena

A ricaniid planthopper
Not described to genus/species.

Undescribed

Undescribed

Undescribed[29]

* Republic

A fulgoroidean hopper
Not described to family/genus/species.

Incertae sedis

†Planophlebia

†Planophlebia gigantea[54]

* Princeton

A hemipteran[32] of uncertain placement

|  Planophlebia gigantea
(1890 illustration)

#### Heteroptera[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Cf. Coreidae

Undescribed

Undescribed[20]

* McAbee

Relatives of leaf-footed bugs
Not described to Family/genus/species

Cydnidae

Undescribed

Undescribed[27]

* Quilchena

A cydnid burrowing bug
Not described to genus/species.

Dinidoridae

Megymenum

Undescribed[27][22]

* Quilchena

A dinidorid shield bug
Not described to genus/species.

Gerridae |

Gerris

Undescribed[20]

* McAbee

gerrine water striders
Not described to genus/species

Limnoporus

†Limnoporus wilsoni[27][55]

* Driftwood

A gerrine water strider[56]

†Telmatrechus |

†Telmatrechus defunctus[27][56]

* Quilchena

A gerrine water strider
First described as "Gerris" defunctus (1910)[43]

|  Telmatrechus defunctus
(1910 illustration)

†Telmatrechus stali[27][56]

* Princeton

A gerrine water strider
First described as "Hygrotrechus" stali (1879),[32]
later moved to "Gerris" stali (1910)[43]

|  Telmatrechus stali
(1890 illustration)

Pentatomidae

Undescribed

Undescribed[24][29]

* Princeton
* Republic

A Shield or stink bug
Not described to genus/species

|  undescribed Pentatomidae

Cf. Pentatomidae

Undescribed

Undescribed[27]

* Quilchena

Pentatomoid shield bugs of uncertain familial placement
Not described to genus/species.

#### Sternorrhyncha[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Aphididae | Undescribed  |

Undescribed[29][27][20]

* McAbee
* Quilchena
* Republic

Aphids
Not described to genus/species.

Undescribed[41]

* Republic

Trace fossils
Woolly aphid leaf rolling on Ulmus leaves
Not described to genus/species

Unidentified

Unidentified

Unidentified

* Driftwood

aphidoid specimens
Not described to genus/species

#### Unidentified[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Unidentified

Unidentified

Unidentified[9]

* Hat Creek

Hemipterans in amber
Not identified in any finer taxonomic detail

### Hymenopterans[edit]

Archibald, Mathewes, & Aase (2023) reported a Titanomyrma species ant queen from Allenby Formation, and noted the range extension for Formiciinae into the highlands, as the subfamily was previously considered a strictly thermophilic ant group. Due to complications arising from preservational distortion during diagenesis, they were unable to determine the correct size of the queen in life. If the distortion was lateral, then compression to bilateral symmetry yielded an adult length of approximately 3.3 cm (1.3 in), placing it the same range as Formicium berryi and F. brodiei, known only from wings, and suggested as possible males. Conversely stretching the fossil to bilateral symmetry results in a larger 5 cm (2.0 in) length estimate, placing it as comparable to queens of T. lubei and T. simillima.[57]

#### Symphyta[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Cephidae |

†Cuspilongus

†Cuspilongus cachecreekensis[4]

* McAbee

A cuspilongine cephid sawfly

Undescribed

"Undescribed[4]

* Horsefly

A cephine cephid sawfly
Not described to genus/species

Cimbicidae[4] |

†Allenbycimbex[58]

†Allenbycimbex morrisae[58]

* Princeton

A cenocimbicine cimbicid sawfly.

†Leptostigma[58] |

†Leptostigma alaemacula[58]

* Republic

A cenocimbicine cimbicid sawfly.

†Leptostigma brevilatum[58]

* McAbee
* Republic

A cenocimbicine cimbicid sawfly.

|  Leptostigma brevilatum?

†Leptostigma fasciatum[58]

* McAbee

A cenocimbicine cimbicid sawfly.

†Leptostigma longiclava[58]

* McAbee

A cenocimbicine cimbicid sawfly.

†Leptostigma longipallidum[58]

* McAbee

A cenocimbicine cimbicid sawfly.

†Leptostigma longitenebricum[58]

* McAbee

A cenocimbicine cimbicid sawfly.

†Leptostigma proxivena[58]

* McAbee

A cenocimbicine cimbicid sawfly.

Unidentified

Unidentified[58]

* McAbee

A cimbicid sawfly of indeterminate subfamily.

Pamphiliidae

†Ulteramus

†Ulteramus republicensis[59]

* Republic

A parasitic wasp

Siricidae |

†Eourocerus

†Eourocerus anguliterreus[60]

* Republic

A siricid horntail.

|  Eourocerus anguliterreus

†Ypresiosirex

†Ypresiosirex orthosemos[4]

* McAbee

A siricine siricid sawfly

Tenthredinidae |

Eriocampa

†Eriocampa tulameenensis[61][4]

* Princeton

An allantine tenthredinid sawfly

|  Eriocampa tulameenensis

Pseudosiobla

†Pseudosiobla campbelli[61][4]

* Horsefly

An allantine tenthredinid sawfly

|  Pseudosiobla campbelli
Undescribed  |

Undescribed[4]

* McAbee

Sawflys of the tenthredinid subfamily Allantinae
Not described

Undescribed[4]

* McAbee

Sawflys of the tenthredinid subfamily Blennocampinae
Not described

Undescribed[4]

* McAbee

Sawflys of the tenthredinid subfamily Nematinae
Not described

Undescribed[4]

* McAbee

Sawflys of the tenthredinid subfamily Tenthredininae
Not described

Undescribed[4]

* Driftwood
* Horsefly
* McAbee
* Quilchena
* Republic

Sawflys of the family Tenthredinidae, unplaced to subfamily
Not described

#### Parasitoida[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Braconidae |

Bracon

Undescribed[32]

* Princeton

A Bracon sensu lato species wasp.

|  Bracon sp.
specimen 69&78
(1890 illustration)

Undescribed

Undescribed[4]

* Driftwood
* Horsefly
* McAbee
* Quilchena
* Republic

braconid parasitic wasps unplaced to subfamily
Not described to genus/species

|  Unidentified Braconidae

Cynipidae

Undescribed

Undescribed[41]

* Princeton
* Republic

Trace fossils
cynipid Cynipoid gallwasp
galling on various host leaves
Not described to genus/species

|  Cynipidae gall on Prunus

Cynipidae (?)

Undescribed

Undescribed[33][4]

* Horsefly
* Quilchena

cynipid Cynipoid gallwasps
Not described to species

Diapriidae

Undescribed

Undescribed[4]

* Driftwood
* Horsefly
* McAbee
* Republic

diapriid diaprioid wasps
Not described to genus/species

Figitidae

Undescribed

Undescribed[4]

* McAbee

figitid cynipoid wasps
Not described to genus/species

Heloridae

Undescribed

Undescribed[4]

* McAbee

helorid proctotrupoid wasps
Not described

Ichneumonidae |

Xorides

†Xorides lambei[43]

* Princeton

A xoridine ichneumon parasitic wasp
First described as Xylonomus lambei.

|  Xorides lambei
(1910 illustration)

Undescribed

Undescribed[33][4]

* Driftwood
* Horsefly
* McAbee
* Quilchena
* Princeton
* Republic

ichneumonid parasitic wasps unplaced to subfamily
Not described

|  Undescribed Ichneumonidae

Megaspilidae

Undescribed

Undescribed[4]

* Hat Creek

ceraphronoid Megaspilid parasitic wasps
Not described

Monomachidae

Undescribed

Undescribed[4]

* McAbee

monomachid diaprioid wasps
Not described

Peradeniidae

Undescribed

Undescribed[4]

* McAbee

peradeniid proctotrupoid wasps
Not described

Proctotrupidae

Undescribed

Undescribed[4]

* Driftwood
* Horsefly
* McAbee
* Republic

proctotrupid parasitic wasps
Not described

Roproniidae

Undescribed

Undescribed[4]

* Republic

roproniid (sensu lato) proctotrupoid wasps
Not described

incertae sedis[4] | Undescribed  |

Undescribed

* Driftwood
* Hat Creek
* McAbee

Chalcidoid superfamily wasps
Not described to family/genus/species

Undescribed[4]

* Hat Creek

mymarommatoid microhymenopterans
Not described to family/genus/species

#### Non-apoidean Aculeates[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Chrysididae

Undescribed

Undescribed[4]

* McAbee

Chrysidoid wasps of the chrysidid subfamily Chrysidinae
Not described to genus/species

Pompilidae

Undescribed

Undescribed[33][4]

* McAbee
* Republic

pompilid spider wasps
Not described

Scoliidae

Undescribed

Undescribed[33][4]

* Princeton
* Republic

Scoliid wasps of the subfamily Archaeoscoliinae
Not described

Trigonalidae

Undescribed

Undescribed[4]

* Falkland
* Quilchena

trigonalid parasitic wasps
Not described

Vespidae

Undescribed

Undescribed[4]

* Driftwood
* McAbee
* Princeton
* Quilchena
* Republic

Vespid wasps
Not described

#### Formicoidea[edit]

subfamily  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Dolichoderinae |

Dolichoderus

Undescribed[4]

* Hat Creek

Dolichoderus species ants.[4] Not described to species

Undescribed

Undescribed[4]

* McAbee

Dolichoderine ants.[4] Not described to genus/species

Formiciinae

†Titanomyrma

Indeterminate[57]

* Princeton

A formiciine titan ant.
Unplaced to species

|  Titianomyrma sp.
Formicinae |

†Eoecophylla

†Eoecophylla quilchenensis[62]

* Quilchena

A weaver ant
related to modern Oecophylla
The most abundant ants at Quilchena

Oecophylla

†Oecophylla kraussei[63]

* Republic

A weaver ant
first described as Camponotites kraussei[64][4][62]

Indeterminate

indeterminate[62]

* McAbee

An weaver ant tribe worker
Possibly belonging to either †Eoecophylla or Oecophylla

Undescribed

Undescribed[4]

* Hat Creek

Formicinae subfamily ants.[4]
Not described to species
First published as Technomyrmex by Poinar et al., (1999)

Myrmeciinae | †Avitomyrmex |

†Avitomyrmex elongatus[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

†Avitomyrmex mastax[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

†Avitomyrmex systenus[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

†Macabeemyrma

†Macabeemyrma ovata[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

†Myrmeciites |

†Myrmeciites? goliath[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

†Myrmeciites? tabanifluviensis[65][4]

* Horsefly

A myrmeciine bulldog ant.[4]

†Myrmeciites herculeanus[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

"Indeterminate"[65][4]

* Falkland
* Republic

bull dog ants
not distinct to species

|  Myrmeciities sp.

†Propalosoma

†Propalosoma gutierrezae[63]

* Republic

A myrmeciine bulldog ant, first described as a rhopalosomatid wasp.[4]

|  Propalosoma gutierrezae
†Ypresiomyrma |

†Ypresiomyrma bartletti[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

†Ypresiomyrma orbiculata[65][4]

* McAbee

A myrmeciine bulldog ant.[4]

|  Ypresiomyrma orbiculata

Undescribed

Undescribed[4]

* McAbee

Myrmeciinae ants.[4] Not described to species

Myrmicine

Undescribed

Undescribed[4]

* Hat Creek

Myrmicine ants of either Leptothorax or Tetramorium[4] Not described to species

Incertae sedis |

†Klondikia

†Klondikia whiteae[66][4]

* Republic

An ant of uncertain subfamily placement[4]

Undescribed

Undescribed[33][4]

* Driftwood
* Hat Creek
* Horsefly
* McAbee
* Princeton
* Quilchena
* Republic

Ants of uncertain subfamily placement.[4]

|  Unidentified formicidae

#### Apoidea[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Angarosphecidae | †Eosphecium |

†Eosphecium naumanni[4]

* Quilchena

A spheciform wasp.

Undescribed[4]

* Driftwood
* McAbee
* Republic

Spheciform wasps.
Not described to species

Undescribed

Undescribed[4]

* McAbee
* Princeton
* Republic

Spheciform wasps, likely not Eosphecium.
Not described to species

Apidae

Undescribed

Undescribed[4]

* McAbee

Apid bees
Not described to species

Halictidae (?)  |

Halictus?

†Halictus? savenyei[4]

* Quilchena

A sweat bee of uncertain generic placement

Undescribed

Undescribed[4]

* McAbee
* Republic

sweat bees
Not described

Megachilidae

Undescribed

Undescribed[4]

* Horsefly
* McAbee
* Republic

Megachilid leaf-cutter bee herbivory trace fossils on leaves
Not described to species

|  Megachilidae damage on Prunus

Sphecidae

Undescribed

Undescribed[4]

* McAbee
* Republic

Sphecid (sensu stricto) wasps
Not described

### Lepidoptera[edit]

A solitary complete adult female lepidopteran fossil has been recovered, but no full descriptive work has been published on the specimen, aside from a single PhD dissertation.[33] Early examination placed the moth in the family Geometridae, but later work has identified it as the oldest member of the tiger moth subfamily Arctiinae.[67] Two additional isolated wing fossils have been found with one tentatively placed within Noctuidae based on the wing venation and structure,[33] while the second has not been placed beyond order level.[68] Laval mining and hole feeding damage on leaves has been attributed to the families Coleophoridae, Gracillariidae, Heliozelidae, Incurvariidae, and Nepticulidae[41][69]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Coleophoridae

Cf. Coleophora

Undescribed[41][68]

* Republic

Trace fossils
Coleophorid hole feeding and larval cases similar to Coleophora
Not described to genus/species

Erebidae

Undescribed

Undescribed[67]

* Republic

An arctiine tiger moth
Not described to genus/species

|  Arctiinae
undescribed

Gracillariidae

Undescribed

Undescribed[41][20]

* McAbee

Trace fossils
Gracillariid leaf mining similar to Phyllocnistinae subfamily mines
Not described to genus/species

Heliozelidae

Undescribed

Undescribed[41][69]

* Republic

Trace fossils
heliozelid leaf mining similar to Antispila mines
Not described to genus/species

Incurvariidae

Aff. Incurvaria

Undescribed[41]

* Republic

Trace fossils
incurvariid leaf mining similar to Incurvaria
Not described to genus/species

Nepticulidae

Stigmella

Undescribed[41]

* Republic

Trace fossils
nepticulid leaf mining referred to Stigmella
Not described to genus/species

Pyralidae?

Stigmella

Undescribed[41]

* Republic

Trace fossils
nepticulid leaf mining referred to Stigmella
Not described to genus/species

Noctuidae

Undescribed

Undescribed[33]

* Quilchena

A possible owlet moth
Not described to genus/species

Undescribed

Undescribed

Undescribed[68]

* Princeton chert

A lepidopteran forewing
Not described to family/genus/species

### Mecopterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Bittacidae

Undescribed

Undescribed[20]

* McAbee

hangingfly specimens
Not described to genus/species

†Cimbrophlebiidae | †Cimbrophlebia |

†Cimbrophlebia brooksi[70]

* Republic

A Cimbrophlebiid scorpionfly

|  Cimbrophlebia brooksi

†Cimbrophlebia flabelliformis[70]

* McAbee

A Cimbrophlebiid scorpionfly

†Cimbrophlebia leahyi[70]

* McAbee

A Cimbrophlebiid scorpionfly

†Cimbrophlebia westae[70]

* Republic

A Cimbrophlebiid scorpionfly

|  Cimbrophlebia westae
†Dinopanorpidae | †Dinokanaga |

†Dinokanaga andersoni[71]

* Republic

A Dinopanorpid scorpion fly

|  Dinokanaga andersoni

†Dinokanaga dowsonae[71]

* Falkland
* Horsefly
* McAbee
* Republic

A Dinopanorpid scorpion fly

|  Dinokanaga dowsonae

†Dinokanaga hillsi[71]

* McAbee

A Dinopanorpid scorpion fly

†Dinokanaga sternbergi[71]

* Republic

A Dinopanorpid scorpion fly

†Dinokanaga webbi[71]

* Horsefly

A Dinopanorpid scorpion fly

†Dinokanaga wilsoni[71]

* Princeton

A Dinopanorpid scorpion fly

Eomeropidae | †Eomerope |

†Eomerope eonearctica[72]

* McAbee

An Eomeropid scorpionfly

†Eomerope macabeensis[73]

* McAbee

An Eomeropid scorpionfly

†Eomerope simpkinsae[72]

* Princeton

An Eomeropid scorpionfly

†Eorpidae | †Eorpa |

†Eorpa elverumi[74]

* Republic

An eorpid scorpionfly

|  Eorpa elverumi

†Eorpa jurgeni[74]

* Quilchena

An eorpid scorpionfly

†Eorpa ypsipeda[74]

* Falkland?
* McAbee
* Republic?

An eorpid scorpionfly

|  Eorpa sp. possibly E. ypsipeda[74]

†Holcorpidae

†Holcorpa

†Holcorpa dillhoffi[75]

* McAbee

A holcorpid scorpionfly

Panorpidae

Undescribed

Undescribed[74]

* McAbee
* Republic

Undescribed common scorpionflies
Not described to genus/species

### Neuropterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Berothidae

†Microberotha

†Microberotha macculloughi[13]

* Hat Creek

A beaded lacewing

Chrysopidae | †Adamsochrysa |

†Adamsochrysa aspera[76]

* McAbee

A nothochrysine green lacewing

†Adamsochrysa wilsoni[76]

* Republic

A nothochrysine green lacewing

|  Adamsochrysa wilsoni
†Archaeochrysa |

†Archaeochrysa profracta[76]

* McAbee

A nothochrysine green lacewing

†Archaeochrysa sanikwa[77]

* Driftwood

A nothochrysine green lacewing

†Lithochrysa

†Lithochrysa borealis[78]

* Driftwood

A nothochrysine green lacewing

†Okanaganochrysa

†Okanaganochrysa coltsunae[76]

* McAbee

A nothochrysine green lacewing

†Protochrysa

†Protochrysa fuscobasalis[76]

* McAbee

A limaiine green lacewing

†Pseudochrysopa

†Pseudochrysopa harveyi[76]

* Driftwood

A nothochrysine green lacewing

†Corydasialidae

†Ypresioneura

†Ypresioneura obscura[79]

* McAbee

A corydasialid lacewing
First suggested to be a megalopterans, later deemed a neuropteran.

Hemerobiidae |

†Archibaldia

†Archibaldia wehri[80][81][82]

* Republic

A hemerobiid lacewing
originally placed in †Cretomerobius
Moved to †Proneuronema (2016)
Moved to †Archibaldia (2023)

Wesmaelius

†Wesmaelius mathewesi[80]

* Quilchena

A hemerobiine brown lacewing.

Undescribed

Undescribed[20]

* McAbee

Brown lacewing specimens
Not described to genus/species

Ithonidae |

†Allorapisma

†Allorapisma chuorum[83]

* Republic

An Ithonid group moth lacewing

|  Allorapisma chuorum
†Palaeopsychops |

†Palaeopsychops dodgeorum[84]

* Quilchena

A polystechotid group moth lacewing
First mentioned by Archibald & Mathewes 2000 as cf Osmylidae or Polystoechotidae

†Palaeopsychops douglasae[84]

* Quilchena

A polystechotid group moth lacewing

†Palaeopsychops marringerae[84]

* Republic

A polystechotid group moth lacewing

|  Palaeopsychops marringerae

†Palaeopsychops setosus[84]

* Horsefly

A polystechotid group moth lacewing

†Palaeopsychops timmi[84]

* Republic

A polystechotid group moth lacewing

|  Palaeopsychops timmi
†Polystoechotites |

†Polystoechotites barksdalae[84]

* Republic

A polystechotid group moth lacewing

|  Polystoechotites barksdalae

†Polystoechotites falcatus[84]

* Republic

A polystechotid group moth lacewing

|  Polystoechotites falcatus

†Polystoechotites lewisi[84]

* Republic

A polystechotid group moth lacewing

|  Polystoechotites lewisi

†Ricaniella

†Ricaniella antiquata[85]

* Princeton

A Polystoechotid-group moth lacewing[86]
moved from Ricania antiquata (1895)[87]

|  Ricaniella antiquata
(1895 illustration)

Undescribed

Undescribed[20]

* McAbee

Polystoechotid-group moth lacewings
Not described to genus/species

Nymphidae |

†Epinesydrion

†Epinesydrion falklandensis[88]

* Falkland

A split-footed lacewing, possibly sister species to †Nymphes georgei

Nymphes

†Nymphes(?) georgei[89]

* Republic

A split-footed lacewing
Possibly related to Epinesydrion

|  Nymphes georgei
Osmylidae | †Osmylidia |

†Osmylidia donnae[90]

* Quilchena

A protosmyline osmylid lacewing

†Osmylidia glastrai[90]

* Republic

A protosmyline osmylid lacewing

Indeterminate[90]

* Driftwood

A protosmyline osmylid lacewing,
not detailed enough for species description

Undescribed

Undescribed[20]

* McAbee

osmylid lacewing specimens
Not described to genus/species

Psychopsidae?

†Ainigmapsychops

†Ainigmapsychops inexspectatus[91]

* Republic

A possible psychopsid lacewing

|  Ainigmapsychops inexspectatus

Undescribed

Undescribed

Undescribed[20]

* McAbee

Members of an undescribed neuropteran family

### Odonata[edit]

Trace fossil evidence of damselflies has been recorded from oviposition scars on various leaves from the Klondike Mountain Formation that have been placed in the ichnogenus Paleoovoidus. Lewis and Carrol (1991) originally identified the damage on an Alnus parvifolia leaf as caused by leaf beetles of the genus Altica. This was later questioned by Conrad Labandeira who noted the scar patterns did not match modern Altica egg laying behaviour.

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Aeshnidae |

†Antiquiala

†Antiquiala snyderae[92]

* Republic

A darner dragonfly

|  Antiquiala snyderae

†Eoshna

†Eoshna thompsonensis[92]

* McAbee

A darner dragonfly

†Idemlinea

†Idemlinea versatilis[92]

* Republic

A darner dragonfly

|  Idemlinea versatilis
†Ypshna |

†Ypshna brownleei[92]

* Republic

A darner dragonfly

|  Ypshna brownleei

†Ypshna latipennata[92]

* McAbee

A darner dragonfly

†Dysagrionidae |

†Dysagrion

†Dysagrion pruettae[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Dysagrion pruettae

†Dysagrionites

†Dysagrionites delinei[93]

* Republic

A dysagrionine cephalozygopteran odonate

|  Dysagrionites delinei
†Okanagrion |

†Okanagrion angustum[93]

* McAbee

A Dysagrionine cephalozygopteran odonate

†Okanagrion beardi[93]

* McAbee

A Dysagrionine cephalozygopteran odonate

†Okanagrion dorrellae[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Okanagrion dorrellae

†Okanagrion hobani[93]

* McAbee
* Republic

A Dysagrionine cephalozygopteran odonate

|  Okanagrion hobani

†Okanagrion liquetoalatum[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Okanagrion liquetoalatum

†Okanagrion lochmum[93]

* McAbee

A Dysagrionine cephalozygopteran odonate

†Okanagrion threadgillae[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Okanagrion threadgillae

†Okanagrion worleyae[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Okanagrion worleyae
†Okanopteryx |

†Okanopteryx fraseri[93]

* McAbee

A Dysagrionine cephalozygopteran odonate

†Okanopteryx jeppesenorum[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Okanopteryx jeppesenorum

†Okanopteryx macabeensis[93]

* McAbee

A Dysagrionine cephalozygopteran odonate

†Stenodiafanus

†Stenodiafanus westersidei[93]

* Republic

A Dysagrionine cephalozygopteran odonate

|  Stenodiafanus westersidei

Cf. †Dysagrionidae

†Allenbya

†Allenbya holmesae[2]

* Princeton

A possible Dysagrionidae odonate.
Not to be confused with the Princeton Chert waterlily Allenbya

|  Allenbya holmesae

Euphaeidae

†Republica

†Republica weatbrooki[94]

* Republic

A gossamerwing damselfly.
Not to be confused Republica,
a plant genus also from Republic

|  Republica weatbrooki

Gomphidae

†Auroradraco

†Auroradraco eos[92]

* McAbee

A club-tailed dragonfly

†Whetwhetaksidae

†Whetwhetaksa

†Whetwhetaksa millerae[93]

* Republic

A possible cephalozygopteran odonate

|  Whetwhetaksa millerae

### Orthoptera[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Acrididae

Unidentified

Unidentified[20]

* McAbee

Catanopine short-horned grasshopper specimens
Not described to genus/species

Gryllacrididae?

Unidentified

Unidentified[20]

* McAbee

Possible leaf-rolling cricket specimens
Not described to genus/species

†Promastacidae

†Promastax

†Promastax archaicus[43]

* Horsefly

A Promastacid grasshopper

|  Promastax archaicus
(1910 illustration)
†Palaeorehniidae |

†Republicopteron

†Republicopteron douseae[95]

* Republic

A palaeorehniid ensiferan.

|  Republicopteron douseae

†Ypopteron

†Ypopteron nicola[95]

* Quilchena

A palaeorehniid ensiferan.
First identified as a prophalangopsine grig[27]

Prophalangopsidae

Unidentified

Unidentified[20]

* McAbee

grig specimens
Not described to genus/species

Tettigoniidae

Unidentified

Unidentified[20]

* McAbee

katydid specimens
Not described to genus/species

### Phasmatodea[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Susumaniidae

†Eoprephasma

†Eoprephasma hichensi[96]

* McAbee
* Republic

A Susumaniine susumanioid stick insect

### Plecoptera[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Unidentified

Unidentified

Unidentified[20]

* McAbee

stonefly specimens
Not described to family/genus/species

### Psocodea[edit]

The only reported Psocodea fossils from the highlands are known from Hat Creek amber. They were mentioned, as "Psocoptera", in passing by Poinar et al. (1999) who did not give any finer taxonomic detail or illustrate any specimens.[9]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Unidentified

Unidentified

Unidentified[9]

* Hat Creek

specimens in amber
Not described to genus/species

### Raphidiopterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Inocelliidae

†Paraksenocellia

†Paraksenocellia borealis[97]

* Driftwood

An inocelliid snakefly

Raphidiidae |

†Archiinocellia

†Archiinocellia oligoneura[98]

* Horsefly

A Raphidiid snakefly

|  Archiinocellia oligoneura
†Megaraphidia |

†Megaraphidia antiquissima[98]

* McAbee

A Raphidiid snakefly

†Megaraphidia hopkinsi[98]

* Princeton

A Raphidiid snakefly

†Megaraphidia klondika[98]

* Republic

A raphidiid snakefly

†Megaraphidia ootsa[98]

* Driftwood

A raphidiid snakefly

### Thrysanoptera[edit]

Poinar et al. (1999) illustrated a Thrips specimen in Hat Creek amber and noted the presence of the order in the fossils they examined, however they did not provide any finer taxonomic details on the affinities of the fossils.[9]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Unidentified

Unidentified

Unidentified[9]

* Hat Creek

specimens in amber
Not described to genus/species

### Trichoptera[edit]

Trichopterans are known mainly from laraval cases and occasional isolated wings.[27]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Limnephilidae

Unidentified

unidentified[29]

* Republic

northern caddisflies
Not described to genus/species

Phryganeidae

Unidentified

Unidentified[29][27]

* Quilchena
* Republic

giant caddisflies
Not described to genus/species

|  Phryganeidae larval case

Unidentified

Unidentified

Unidentified[24][27][20]

* Driftwood
* Horsefly
* McAbee
* Quilchena

Isolated wings and larval cases
Not described to genus/species

## Vertebrates[edit]

### "Fish"[edit]

The first fish to be described from the Okanagan Highlands were recovered from Allenby Formation shales and subsequently studied by Edward Drinker Cope who named Amyzon brevipinne in 1894. The next descriptive work for a fish came in 1916 with the naming of "Lucious" rosei by Louis Hussakof from Tranquille Formation fossils collected at "Red point" on Kamloops Lake in 1914.[99] "Lucious" rosei was redescribed in 1966 by Ted Cavander, who moved the species to a new genus Eohiodon placed into the mooneye family Hiodontidae. The largest body of work for fish of the Highlands was by Mark Wilson (1977) who published a monograph detailing the Canadian highlands formations fish fauna, naming four new species in three new genera, plus redescribing both "Amyzon" brevipinne and "Eohiodon" rosei. The monograph added the families Salmonidae with Eosalmo driftwoodensis, Libotoniidae with Libotonius blakeburnensis, and Moronidae with Priscacara aquilonia.[24] A year later the first species from the Republic area, "Eohiodon" woodruffi was described by Wilson,[100] and the second Republic species Libotonius pearsoni followed in 1979.[101] in 1982 the final new fish species named from the highlands , Amia hesperia, was described, being initially placed by Wilson in the modern bowfin genus Amia.[102] This placement was later questioned by Lance Grande and William Bemis (1998), who noted that due to preservational orientation of the A. hesperia holotype, generic placement of the species was problematic. Phylogenetic analysis of Amiidae fossils by Grande and Bemis found the fossil as a member of the amiinae subfamily, but with key mouth anatomy missing, were unable to determine if Amia or the extinct genus Cyclurus was correct.[103] In 2021 fossils of "Amyzon" brevipinne were redescribed by Juan Liu based on the holotype and additional fossils from the Allenby Formation, and based on the anatomical differences between the species and the type species of Amyzon mentale determined that the Princeton fossils were part of a different genus. As such Liu moved the species to the new genus Wilsonium.[104]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Amiidae | Amia?  |

†Amia? hesperia[102]

* Princeton

A bowfin of uncertain genus placement[103]

†Amia? hesperia?[28][102]

* Driftwood
* Princeton
* Quilchena
* Republic

isolated bowfin scales[103]

|  Amia? scale
Catostomidae | †Amyzon |

†Amyzon aggregatum[24][105]

* Horsefly
* Republic?

A sucker

Unidentified[24][105][104][20]

* Driftwood
* McAbee
* Republic

A sucker
Republic specimens formerly identified as A. aggregatum[24]

|  Amyzon sp.
†Wilsonium |

†Wilsonium brevipinne[24][106][104]

* Princeton

A catostomid sucker,
first described as Amyzon brevipinne (1894)
Moved to Wilsonium brevipinne in 2021

Cf. †Wilsonium brevipinne[17][28][104]

* Quilchena

A catostomid sucker,
Not described to species
similar to Wilsonium brevipinne

Hiodontidae | Hiodon |

†Hiodon rosei[24][100][107][20]

* Horsefly
* Kamloops Lake
* McAbee
* Princeton

A mooneye, first described as Lucious rosei (1916),
Moved to Eohiodon rosei in 1966,
Moved to Hiodon rosei in 2008[100][107]

†Hiodon woodruffi[100][107]

* Horsefly
* Republic

A mooneye, first described as Eohiodon woodruffi.[100]
Moved to Hiodon woodruffi in 2008[107]

|  Hiodon woodruffi

Unidentified[28]

* Quilchena

A mooneye jaw.
Not identified to species

†Libotoniidae | †Libotonius |

†Libotonius blakeburnensis[24]

* Princeton

A sand roller relative.

†Libotonius pearsoni[101]

* Republic

A sand roller relative.

|  Libotonius pearsoni

Moronidae

†Priscacara

†Priscacara aquilonia[24]

* Horsefly

A temperate bass

Salmonidae | †Eosalmo |

†Eosalmo driftwoodensis[24][108][20]

* Driftwood Canyon
* McAbee
* Princeton
* Republic

A basal Salmon

|  Eosalmo driftwoodensis

Cf. †Eosalmo driftwoodensis[17][28]

* Quilchena

A basal Salmon
Not described to species
Similar to E. driftwoodensis

### Reptiles[edit]

The only reptile fossils known from the Okanagan highlands come from the Allenby Formation. A soft-shelled turtle is known from the "Ashnola shales" unit and unidentified turtle bone are known from the interbedded Princeton Chert. The soft shelled turtle was first discovered by James Basinger from dark shale layers above the chert and reported by Wilson (1982).[102] The unidentified turtle bones were found preserved within the chert layers and first reported by Stockey and Pigg (1994).[109] In his 1995 Masters thesis, G. Guthrie listed an isolated tooth from the Quilchena site as from a crocodile, which would have been the only instance of a crocodylian in the highlands.[28] The taxonomic affinity was later revised after further examination and Mathewes et al. (2016) listed the specimen as an unidentified fish tooth.[110]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Trionychidae

Cf. Apalone or Aspideretes

undescribed[102][111][22][20]

* Princeton

A soft shelled turtle
Found in the Ashnola shales over the chert

Undescribed

Undescribed

undescribed[109]

* Princeton chert

A turtle,
Bones preserved in the Princeton Chert

### Birds[edit]

A small avifauna is known from the Okanagan Highlands, but due to the incomplete nature of the fossils, placement of studied specimens has been tentative at best.[112] Gerald Mayr et al. (2019) published an initial overview of the fossils with descriptions and commentary of the material, noting the taxa identified were all previously unknown to Northwestern North American Eocene sites. Despite the tentative nature of the fossil identifications, the Highlands sites are the richest Paleogene avifauna described from Canada. Mayr et al. (2019) posited that the fossils likely represent the more common species in the avifauna of the Highlands, but at the same time, include taxa that are considered rare or absent in the most studied avifaunas from the same time frame.[112] Additional evidence of birds at some sites consists of preserved egested bird pellets, which are composed of randomly grouped fish bone clumps, occasionally including multiple fish or insects.[28]

Isolated feathers are also known from several of the sites and have not described in detail.[112]

Order/Clade  | Family  | Material  | Sites  | Notes  | Images

|
|
|
|
|

Cf. Coliiformes[112]

indeterminate

partial skeleton
skull, left wing, and right wing portions

* Driftwood

a large possible Coliiform,
too incomplete for firm identification

Cf. Gaviiformes[112]

indeterminate

left wing and partial right wing, with feathers

* McAbee

a possible gaviiform,
too incomplete for firm identification

Gruiformes[112]

Cf. †Songziidae

articulated postcranial skeleton

* Driftwood

A possible songziid bird,
lacking specific morphologic characters for a firm placement.

Psittacopasserae[112]

Cf. †Zygodactylidae

complete, but poorly preserved, skeleton

* McAbee

a possible zygodactylid,
poor preservation of the foot prevented firm identification

indeterminate[112]

indeterminate

partial skeletons

* McAbee
* Princeton
* Republic

partial skeletons missing details for identification.
Republic specimen shows unique cranial pitting.

|  Indeterminate skeleton
McAbee site

indeterminate[17][28][20][112]

indeterminate

feathers

* Horsefly
* McAbee
* Quilchena
* Republic

isolated feathers

|  unidentified feather, Republic

### Mammals[edit]

Eocene mammals are exclusively known from sites in, or possibly in, the Okanagan Highlands. The earliest reported mammals were of teeth from the Princeton area in 1935, with one of the fossils subsequently being "lost". More recent work in 2014 and 2017 on fossils from Driftwood and Princeton have expanded the mammal families to three, possibly four, and an undescribed Quilchena fossil being identified as a "lipotyphla".[113][5] The record of Brontotheriidae is uncertain due to the split opinion regarding inclusion of the Quesnel area sediments as part of the Highlands.[5][4][2]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Erinaceidae |

†Silvacola

†Silvacola acares[113]

* Driftwood

A hedgehog and moonrat relative

†Entomolestes

undescribed[113][5]

* Princeton

An unidentified hedgehog relative,
specimen possibly lost.

Esthonychidae | †Trogosus |

†Trogosus latidens[114][102][5]

* Princeton

A tillodont species

†Trogosus sp.[5]

* Princeton

An indeterminate tillodont
smaller than T. latidens

†Helaletidae

Cf. †Heptodon

indeterminate[113]

* Driftwood

A helaletid tapir relative.

undescribed

undescribed

undescribed[5]

* Quilchena

A "lipotyphlan"

## Trace fossils[edit]

Pellets of fish bone and other animal material which were likely eaten by larger predators and then regurgitated have been reported from the Quilchena and Republic sites. These traces, called regurgitalites, have so far been understudied, with only a few passing mentions in Okanagan highlands literature.[112]

## Quesnelian fauna[edit]

If the Quesnel sites are included as part of the Greater Okanagan Highlands per Archibald et al. (2018) the fauna of the region is expanded by a number of insect taxa, an additional arachnid, and a brontothere.

### Arachnids[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Araneae

†Araneaovoius

†Araneaovoius columbiae[115]

* Quesnel

An orb-web spider egg sack.
First described as "Aranea" columbiae (1878)[116]

|  Araneaovoius columbiae
(1890 illustration

### Coleopterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Nitidulidae

Prometopia

†Prometopia depilis[117]

* Quesnel

A sap beetle

|  Prometopia depilis
(1890 illustration)

### Dipterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Anthomyiidae

Anthomyia

†Anthomyia burgessi[117][49]

* Quesnel

An anthomyiid fly.
Considered nomen dubium without discussion by Michelsen (1996).[118]

|  Anthomyia burgessi
(1890 illustration)

†Anthomyia inanimata[117]

* Quesnel

An anthomyiid fly.
Considered nomen dubium without discussion by Michelsen (1996).[118]

|  Anthomyia inanimata
(1890 illustration)

Asilidae

Undescribed

Undescribed[32]

* Quesnel

An undescribed robber fly

Dolichopodidae

Dolichopus

Undescribed[117]

* Quesnel

An undescribed long-legged fly.

|  Dolichopus sp.
specimen 14651
(1890 illustration)

Heleomyzidae

Heteromyza

†Heteromyza senilis[117]

* Quesnel

A heleomyzid fly.

|  Heteromyza senilis
(1890 illustration)

Lauxaniidae

"Lonchaea"

†"Lonchaea" senescens[42]

* Quesnel

A lauxaniid fly. First placed in Lonchaea[117]
Referred to Lauxaniidae without redescription.(1994)

|  "Lonchaea" senescens
(1890 illustration)

Mycetophilidae

Boletina

†Boletina sepulta[117]

* Quesnel

A fungus gnat.

|  Boletina sepulta
(1890 illustration)

Brachypeza

†Brachypeza abita[117]

* Quesnel

A mycetophiline fungus gnat.

|  Brachypeza abita
(1890 illustration)

†Brachypeza procera[117]

* Quesnel

A mycetophiline fungus gnat.

|  Brachypeza procera
(1890 illustration)

Trichonta

†Trichonta dawsoni[117]

* Quesnel

A mycetophiline fungus gnat.

|  Trichonta dawsoni
(1890 illustration)

Pallopteridae

Palloptera

†Palloptera morticina[117]

* Quesnel

A flutter-wing fly.

|  Palloptera morticina
(1890 illustration)

Sciaridae

Sciara

†Sciara deperdita[116]

* Quesnel

A sciarine dark-winged fungus gnat.

|  Sciara deperdita
(1890 illustration)

Sciomyzidae

Sciomyza

†Sciomyza revelata[117]

* Quesnel

A marsh fly.

|  Sciomyza revelata
(1890 illustration)

Ulidiidae

†Lithortalis

†Lithortalis picta[117]

* Quesnel

A picture-winged fly.

|  Lithortalis picta
(1890 illustration)

### Hemipterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Pentatomidae

†Teleoschistus

†Teleoschistus antiquus[49]

* Quesnel

A shield bug
first described as "Euschistus" antiquus (1878).[116]

|  Teleoschistus antiquus
(1890 illustration)

Incertae sedis

†Geranchon

†Geranchon petrorum[49]

* Quesnel

An aphidomorph of uncertain placement
First described as "Lachnus" petrorum (1877)[117]

|  Geranchon petrorum
(1890 illustration)

incertae sedis

†Sbenaphis

†Sbenaphis quesneli[49]

* Quesnel

An aphidoid of uncertain placement
First described as "Lachnus" quesneli[116]

|  Sbenaphis quesneli
(1890 illustration)

### Hymenopterans[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Formicidae

Aphaenogaster

†Aphaenogaster longaeva[117]

* Quesnel

A myrmicine ant, possibly nomen dubium.[119]

|  Aphaenogaster longaeva
(1890 illustration)

†Calyptites

†Calyptites antediluvianum[117]

* Quesnel

An ant of uncertain placement.

|  Calyptites antediluvianum
(1890 illustration)

Dolichoderus

†Dolichoderus obliteratus[120]

* Quesnel

A dolichoderine ant
First described as "Hypoclinea" obliterata[117]

|  Dolichoderus obliteratus
(1890 illustration)

Formica

†Formica arcana[117]

* Quesnel

A formicine ant

|  Formica arcana
(1890 illustration)

Ichneumonidae

Pimpla

†Pimpla decessa[117]

* Quesnel

A pimpline ichneumon parasitic wasp

|  Pimpla decessa
(1890 illustration)

†Pimpla saxea[117]

* Quesnel

A pimpline ichneumon parasitic wasp

|  Pimpla saxea
(1890 illustration)

†Pimpla senecta[117]

* Quesnel

A pimpline ichneumon parasitic wasp

|  Pimpla senecta
(1890 illustration)

### Neuroptera[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Hemerobiidae

†Bothromicromus

†Bothromicromus lachlani[116]

* Quesnel

A hemerobiid lacewing of uncertain subfamily placement

|  Bothromicromus lachlani
(1890 illustration)

### Mammals[edit]

Family  | Genus  | species  | Sites  | Notes  | Images

|
|
|
|
|

Brontotheriidae

indeterminate

indeterminate[121]

* Quesnel

A Brontotheriina subtribe Brontothere
Not identifiable to genus
Known from isolated teeth

## References[edit]

1. ^ a b c Moss, P.; Greenwood, D.; Archibald, S. (2005). "Regional and local vegetation community dynamics of the Eocene Okanagan Highlands (British Columbia – Washington State) from palynology". Canadian Journal of Earth Sciences. 42 (2): 187–204. Bibcode:2005CaJES..42..187M. doi:10.1139/E04-095.
2. ^ a b c d Archibald, S. B.; Cannings, R. A. (2022). "The first Odonata from the early Eocene Allenby Formation of the Okanagan Highlands, British Columbia, Canada (Anisoptera, Aeshnidae and cf. Cephalozygoptera, Dysagrionidae)". The Canadian Entomologist. 154 (1): e29. doi:10.4039/tce.2022.16. S2CID 250035713.
3. ^ Höy, T.; Friedman, R.; Gabites, J. Paleogene Penticton Group, Boundary area, Southern British Columbia (Parts of NTS 082E): Geochronology and Implications for Precious Metal Mineralization (PDF) (Report). Geoscience BC Summary of Activities 2020: Minerals, Geoscience BC, Report 2021-01. Geoscience BC. pp. 55–66.
4. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj bk bl bm bn bo bp bq br bs bt bu bv bw bx Archibald, S. B.; Rasnitsyn, A. P.; Brothers, D. J.; Mathewes, R. W. (2018). "Modernisation of the Hymenoptera: ants, bees, wasps, and sawflies of the early Eocene Okanagan Highlands of western North America". The Canadian Entomologist. 150 (2): 205–257. doi:10.4039/tce.2017.59. ISSN 0008-347X. S2CID 90017208.
5. ^ a b c d e f g Eberle, J.J.; Greenwood, D.R. (2017). "An Eocene brontothere and tillodonts (Mammalia) from British Columbia, and their paleoenvironments". Canadian Journal of Earth Sciences. 54 (9): 981–992. Bibcode:2017CaJES..54..981E. doi:10.1139/cjes-2017-0061. hdl:1807/77901.
6. ^ a b c Archibald, S.; Greenwood, D.; Smith, R.; Mathewes, R.; Basinger, J. (2011). "Great Canadian Lagerstätten 1\. Early Eocene Lagerstätten of the Okanagan Highlands (British Columbia and Washington State)". Geoscience Canada. 38 (4): 155–164.
7. ^ Feldmann, R.; Schweitzer, C.; Leahy, J. (2011). "New Eocene crayfish from the McAbee Beds in British Columbia: First record of Parastacoidea in the Northern Hemisphere". Journal of Crustacean Biology. 31 (2): 320–331. doi:10.1651/10-3399.1.
8. ^ a b Archibald, S. B.; Makarkin, V. N. (2006). "Tertiary giant lacewings (Neuroptera: Polystoechotidae): revision and description of new taxa from western North America and Denmark" (PDF). Journal of Systematic Palaeontology. 4 (2): 119–155. doi:10.1017/S1477201906001817. S2CID 55970660.
9. ^ a b c d e f g h i j k l m n o p q Poinar, G.; Archibald, S.; Brown, A. (1999). "New amber deposit provides evidence of early Paleogene extinctions, paleoclimates, and past distributions". The Canadian Entomologist. 131 (2): 171–177. doi:10.4039/Ent131171-2. S2CID 85718312.
10. ^ a b Grimaldi, D.; Sunderlin, D.; Aaroe, G.; Dempsky, M.; Parker, N.; Tillery, G.; White, J.; Barden, P.; Nascimbene, P.; Williams, C. (2018). "Biological inclusions in amber from the Paleogene Chickaloon Formation of Alaska". American Museum Novitates (3908): 1–37.
11. ^ Pigg, K. B.; DeVore, M. L. (2016). "A review of the plants of the Princeton chert (Eocene, British Columbia, Canada)". Botany. 94 (9): 661–681. doi:10.1139/cjb-2016-0079. hdl:1807/73571.
12. ^ a b c McKellar, R. C.; Wolfe, A. P.; Penney, D. (2010). "Chapter 9: Canadian amber". In Penney, D. (ed.). Biodiversity of Fossils in Amber from the Major World Deposits. Siri Scientific Press. pp. 149–166. ISBN 978-0-9558636-4-6.
13. ^ a b c Archibald, S.B.; Makarkin, V.N. (2004). "New genus of minute Berothidae (Neuroptera) from Early Eocene amber of British Columbia". The Canadian Entomologist. 136 (1): 61–76. CiteSeerX 10.1.1.552.2285. doi:10.4039/n03-043. S2CID 36459014.
14. ^ a b c d Wilson, M. (1977). "New records of insect families from the freshwater Middle Eocene of British Columbia". Canadian Journal of Earth Sciences. 14 (5): 1139–1155. Bibcode:1977CaJES..14.1139W. doi:10.1139/e77-104.
15. ^ Wilson, M. (1978). "Paleogene insect faunas of western North America". Quaestiones Entomologicae. 14 (1): 13–34.
16. ^ a b c d e f g h Russell, L. S. (1957). "Mollusca from the Tertiary of Princeton, British Columbia". National Museum of Canada Bulletin. 147: 84–95.
17. ^ a b c d e Wilson, M. (1987). "Predation as a source of fish fossils in Eocene lake sediments". PALAIOS. 2 (5): 497–500. Bibcode:1987Palai...2..497W. doi:10.2307/3514620. JSTOR 3514620.
18. ^ a b Pigg, K. B.; Bryan, F. A.; DeVore, M. L. (2018). "Paleoallium billgenseli gen. et sp. nov.: fossil monocot remains from the latest Early Eocene Republic Flora, northeastern Washington State, USA". International Journal of Plant Sciences. 179 (6): 477–486. doi:10.1086/697898. S2CID 91055581.
19. ^ a b Selden, P.A.; Penney, D. (2009). "A fossil spider (Araneae: Pisauridae) of Eocene age from Horsefly, British Columbia, Canada". Contributions to Natural History. 12: 1269–1282.
20. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as Wilson, M.V.H. 2009. McAbee Fossil Site Assessment Report. 60 pp.Online PDF. Accessed 17 May 2021.
21. ^ a b Wehr, W. "Middle Eocene insects and plants of the Okanogan Highlands". In Martin, J. (ed.). Contributions to the Paleontology and Geology of the West Coast. Seattle, Washington: University of Washington Press. pp. 99–109.
22. ^ a b c d e f g h i Greenwood, D.; Archibald, S.; Mathewes, R.; Moss, P. (2005). "Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape". Canadian Journal of Earth Sciences. 42 (2): 167–185. Bibcode:2005CaJES..42..167G. doi:10.1139/e04-100.
23. ^ a b Labandeira, C.C. (2021). "Ecology and evolution of gall-inducing arthropods: The pattern from the terrestrial fossil record". Frontiers in Ecology and Evolution. 9. doi:10.3389/fevo.2021.632449.
24. ^ a b c d e f g h i j k l m n o p q r s t u Wilson, MVH (1977). "Middle Eocene freshwater fishes from British Columbia". Life Sciences Contributions, Royal Ontario Museum. 113: 1–66.
25. ^ a b c Wehr, W. C.; Barksdale, L. L. (1995). "Implications of middle Eocene feathers and crayfish from Republic, Washington". Washington Geology. 23 (4): 6–10.
26. ^ a b c Feldmann, R.; Schweitzer, C.; Leahy, J. (2011). "New Eocene crayfish from the McAbee Beds in British Columbia: First record of Parastacoidea in the Northern Hemisphere". Journal of Crustacean Biology. 31 (2): 320–331. doi:10.1651/10-3399.1.
27. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad Archibald, S.B.; Mathewes, R.W. (2000). "Early Eocene insects from Quilchena, British Columbia, and their paleoclimatic implications" (PDF). Canadian Journal of Zoology. 78 (8): 1441–1462. doi:10.1139/cjz-78-8-1441.
28. ^ a b c d e f g h Guthrie, G. H. (1995). A high resolution palaeoecological analysis of an Eocene fossil locality from Quilchena, British Columbia (MA thesis). Simon Fraser University.
29. ^ a b c d e f g h i j k l m n o p q r s t u v w Lewis, S.E. (1992). "Insects of the Klondike Mountain Formation, Republic, Washington". Washington Geology. 20 (3): 15–19.
30. ^ a b c Lawrence, J.F.; Archibald, S.B.; Ślipiński, A. (2008). "A new species of Prionoceridae (Coleoptera: Cleroidea) from the Eocene of British Columbia, Canada". Annales Zoologici. 58 (4): 689–693. doi:10.3161/000345408X396620. S2CID 55873252.
31. ^ Bousquet, Y (2012). "Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico". ZooKeys (245): 1570. doi:10.3897/zookeys.245.3416. PMC 3577090. PMID 23431087.
32. ^ a b c d e f g h i j k l m n o p q Scudder, S. H (1879). "Appendix A. The fossil insects collected in 1877, by Mr. G.M. Dawson, in the interior of British Columbia". Geological Survey of Canada, Report of Progress for. 1877–1878: 175–185.
33. ^ a b c d e f g h i j k l m n o p q r s t u v w Douglas, S.; Stockey, R. (1996). "Insect fossils in middle Eocene deposits from British Columbia and Washington State: faunal diversity and geological range extensions". Canadian Journal of Zoology. 74 (6): 1140–1157.
34. ^ Legalov, A. A. (2013). "New and little known weevils (Coleoptera: Curculionoidea) from the Paleogene and Neogene". Historical Biology: An International Journal of Paleobiology. 25 (1): 59–80. doi:10.1080/08912963.2012.692681. S2CID 86584002.
35. ^ Legalov, A. A. (2015). "Fossil Mesozoic and Cenozoic weevils (Coleoptera, Obrienioidea, Curculionoidea)". Paleontological Journal. 49 (13): 1442–1513. doi:10.1134/S0031030115130067. S2CID 87912009.
36. ^ a b c d Scudder, S. H (1895). "Canadian fossil insects, myriapods and arachnids, Vol II. The Coleoptera hitherto found fossil in Canada". Geological Survey of Canada Contributions to Canadian Palaeontology. 2: 5–26.
37. ^ Archibald, S.B.; Morse, G.; Greenwood, D.R.; Mathewes, R.W. (2014). "Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum". Proceedings of the National Academy of Sciences. 111 (22): 8095–8100. Bibcode:2014PNAS..111.8095A. doi:10.1073/pnas.1323269111. PMC 4050627. PMID 24821798.
38. ^ a b Kundrata, R.; Packova, G.; Prosvirov, A.; Hoffmannova, J. (2021). "The fossil record of Elateridae (Coleoptera: Elateroidea): Described species, current problems and future prospects". Insects. 12 (286): 20. doi:10.3390/insects12040286. PMC 8064311. PMID 33805978.
39. ^ a b c Lewis, S.E. (1994). "Fossil earwigs (Dermaptera) from the Klondike Mountain Formation (middle Eocene) of Republic, Washington". Washington Geology. 22 (1): 39–40.
40. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw Rice, H. M. A (1959). "Fossil Bibionidae (Diptera) from British Columbia". Geological Survey of Canada Bulletin. 55: 1–36.
41. ^ a b c d e f g h i j k Labandeira, C. C. (2002). "Paleobiology of middle Eocene plant-insect associations from the Pacific Northwest: a preliminary report". Rocky Mountain Geology. 37 (1): 31–59. doi:10.2113/gsrocky.37.1.31.
42. ^ a b c Evenhuis (1994). Catalogue of the Fossil Flies of the World (Insecta: Diptera). Backhuys Publishers. pp. 1–600.
43. ^ a b c d e f g h i j k l m n o p q r s t Handlirsch, A. (1910). "Canadian fossil Insects. 5. Insects from the Tertiary lake deposits of the southern interior of British Columbia, collected by Mr. Lawrence M. Lambe". Contributions to Canadian Palaeontology. 2 (3): 93–129.
44. ^ a b Archibald, SB; Kehlmaier, C; Mathewes, RW (2014). "Early Eocene big headed flies (Diptera: Pipunculidae) from the Okanagan Highlands, western North America". The Canadian Entomologist. 146 (4): 429–443. doi:10.4039/tce.2013.79. S2CID 55738600.
45. ^ a b c d e f g h i 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". Palaeogeography, Palaeoclimatology, Palaeoecology. 371: 1–8. Bibcode:2013PPP...371....1A. doi:10.1016/j.palaeo.2012.10.043.
46. ^ Gentilini, G. (1991). "Bibionidae (Diptera: Nematocera) from the Upper Miocene of Monte Castellaro (Marche, Central Italy)". Bolletino del Museo Civico di Storia Naturale, Verona. 15: 425–446.
47. ^ Scuddar, S. H. (1892). "Some Insects of special interest from Florissant, Colorado and other points in the Tertiaries of Colorado and Utah". Bulletin of the United States Geological Survey. 93: 1–25.
48. ^ Fitzgerald, S. J. (1999). "A new species of Plecia from the Green River Formation and new combinations of fossil Bibionidae (Diptera)". Great Basin Naturalist. 59: 182–187. doi:10.5962/bhl.part.15821.
49. ^ a b c d e f g Scudder, S. H. (1890). "The Tertiary insects of North America". United States Geological Survey of the Territories, Washington: 615. ISBN 9780665263149.
50. ^ Handlirsch, A. (1909). "Zur Phylogenie und Flügelmorphologie der Ptychopteriden (Dipteren)". Annalen des Kaiserlich-Königlichen Naturhistorischen Hofmuseums. 23: 263–272.
51. ^ Lewis, S. E.; Wehr, W. (1993). "Fossil mayflies from Republic, Washington" (PDF). Washington Geology. 21 (1): 35–37.
52. ^ a b c Sinitchenkova, N. D. (1999). "A new mayfly species of the extant genus Neoephemera from the Eocene of North America (Insecta: Ephemerida=Ephemeroptera)" (PDF). Paleontological Journal. 33 (4): 403–405. Archived from the original (PDF) on 2011-09-30.
53. ^ a b c d e f g h i Scudder, S. H (1895). "Canadian fossil insects, myriapods and arachnids, Vol I. The Tertiary Hemiptera of British Columbia". Geological Survey of Canada Contributions to Canadian Palaeontology. 2: 5–26.
54. ^ Szwedo, J.; Lefebvre, F. (2004). Fossil planthoppers of the world (Hemiptera: Fulgoromorpha). An annotated catalogue with notes on Hemiptera classification. Warszawa: Studio, 1. pp. 48–49. ISBN 83-912236-8-X.
55. ^ Andersen, N.M. (1998). "Water striders from the Paleogene of Denmark and review of fossil record and evolution of semi aquatic bugs (Hemiptera Gerromorpha)". Biologiske Skrifter. 50: 1–157.
56. ^ a b c Damgaard, J. (2008). "Evolution of the semi-aquatic bugs (Hemiptera: Heteroptera: Gerromorpha) with a re-interpretation of the fossil record". Acta Entomologica Musei Nationalis Pragae. 48: 251–268.
57. ^ a b Archibald, S.; Mathewes, R.; Aase, A. (2023). "Eocene giant ants, Arctic intercontinental dispersal, and hyperthermals revisited: discovery of fossil Titanomyrma (Hymenoptera: Formicidae: Formiciinae) in the cool uplands of British Columbia, Canada". The Canadian Entomologist. 155 (e6).
58. ^ a b c d e f g h i j k Archibald, S. B.; Rasnitsyn, A. P. (2023). "Cimbicidae (Hymenoptera, "Symphyta") in the Paleogene: revision, the new subfamily Cenocimbicinae, and new taxa from the Eocene Okanagan Highlands". Zootaxa. 5278 (1): 1–38. doi:10.11646/zootaxa.5278.1.1.
59. ^ Archibald, S.B.; Rasnitsyn, A.P. (2015). "New early Eocene Siricomorpha (Hymenoptera: Symphyta: Pamphiliidae, Siricidae, Cephidae) from the Okanagan Highlands, western North America". The Canadian Entomologist. 148 (2): 209–228. doi:10.4039/tce.2015.55. S2CID 85743832.
60. ^ Archibald, S. B.; Rasnitsyn, A. P. (2022). "The early Eocene Eourocerus anguliterreus gen. et sp. nov (Hymenoptera, Siricidae) from Republic, Washington". Zootaxa. 5105 (2): 289–295. doi:10.11646/zootaxa.5105.2.8. PMID 35391300. S2CID 247240762.
61. ^ a b Rice, H.M.A. (1968). "Two Tertiary sawflies, (Hymenoptera - Tenthredinidae), from British Columbia". Geological Survey of Canada. 67 (59): 1–21.
62. ^ a b c Archibald, S. B.; Mathewes, R. W.; Perfilieva, K. S. (2024). "Fossil weaver ants (Hymenoptera, Formicidae, Oecophyllini) of the early Eocene Okanagan Highlands of far-western North America". The Canadian Entomologist. 156. e2. doi:10.4039/tce.2023.27.
63. ^ a b Dlussky, G. M.; Rasnitsyn, A. P. (1999). "Two new species of aculeate hymenopterans (Vespida=Hymenoptera) from the Middle Eocene of the United States". Paleontological Journal. 33: 546–549.
64. ^ Perfilieva, K. S.; Dubovikoff, D. A.; Dlussky, G. M. (2017). "Miocene ants (Hymenoptera, Formicidae) from Crimea". Paleontological Journal. 51 (4): 391–401. doi:10.1134/S0031030117040098. S2CID 90536477.
65. ^ a b c d e f g h i j Archibald, S.B.; Cover, S. P.; Moreau, C. S. (2006). "Bulldog Ants of the Eocene Okanagan Highlands and History of the Subfamily (Hymenoptera: Formicidae: Myrmeciinae)" (PDF). Annals of the Entomological Society of America. 99 (3): 487–523. doi:10.1603/0013-8746(2006)99[487:BAOTEO]2.0.CO;2.
66. ^ Dlussky, G. M.; Rasnitsyn, A. P. (2003). "Ants (Hymenoptera: Formicidae) of Formation Green River and some other Middle Eocene deposits of North America". Russian Entomological Journal. 11 (4): 411–436.
67. ^ a b Toussaint, E. F.; Condamine, F. L.; Kergoat, G. J.; Capdevielle-Dulac, C.; Barbut, J.; Silvain, J. F.; Le Ru, B. P. (2012). "Palaeoenvironmental shifts drove the adaptive radiation of a noctuid stemborer tribe (Lepidoptera, Noctuidae, Apameini) in the Miocene". PLOS ONE. 7 (7): e41377. Bibcode:2012PLoSO...741377T. doi:10.1371/journal.pone.0041377. PMC 3409182. PMID 22859979.
68. ^ a b c Sohn, Jae-Cheon; Labandeira, Conrad; Davis, Donald; Mitter, Charles (30 April 2012). "An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world". Zootaxa. 3286 (1): 59. doi:10.11646/zootaxa.3286.1.1.
69. ^ a b Grimaldi, D.; Engel, M. S. (2005). Evolution of the Insects. Cambridge University Press. ISBN 978-0-521-82149-0.
70. ^ a b c d Archibald, S. B. (2009). "New Cimbrophlebiidae (Insecta: Mecoptera) from the Early Eocene at McAbee, British Columbia, Canada and Republic, Washington, USA" (PDF). Zootaxa. 2249: 51–62. doi:10.11646/zootaxa.2249.1.5. S2CID 16035433.
71. ^ a b c d e f Archibald, S.B. (2005). "New Dinopanorpidae (Insecta: Mecoptera) from the Eocene Okanagan Highlands (British Columbia, Canada and Washington State, USA)". Canadian Journal of Earth Sciences. 42 (2): 119–136. Bibcode:2005CaJES..42..119A. doi:10.1139/e04-073.
72. ^ a b Archibald, S. B.; Rasnitsyn, A. P. (2018). "Two new species of fossil Eomerope (Mecoptera: Eomeropidae) from the Ypresian Okanagan Highlands, far-western North America, and Eocene Holarctic dispersal of the genus". The Canadian Entomologist. 150 (3): 393–403. doi:10.4039/tce.2018.13. S2CID 90119028.
73. ^ Archibald, S.B.; Rasnitsyn, A.P.; Akhmetiev, M.A. (2005). "The ecology and distribution of Cenozoic Eomeropidae (Mecoptera), and a new species of Eomerope Cockerell from the Early Eocene McAbee locality, British Columbia, Canada". Annals of the Entomological Society of America. 98 (4): 503–514. doi:10.1603/0013-8746(2005)098[0503:EADOCE]2.0.CO;2.
74. ^ a b c d e Archibald, SB; Mathewes, RW; Greenwood, DR (2013). "The Eocene apex of panorpoid scorpionfly family diversity". Journal of Paleontology. 87 (4): 677–695. doi:10.1666/12-129. S2CID 88292018.
75. ^ Archibald, SB (2010). "Revision of the scorpionfly family Holcorpidae (Mecoptera), with description of a new species from Early Eocene McAbee, British Columbia, Canada" (PDF). Annales de la Société Entomologique de France. 46 (1–2): 173–182. doi:10.1080/00379271.2010.10697654.
76. ^ a b c d e f Vladimir N. Makarkin; S. Bruce Archibald (2013). "A Diverse New Assemblage of Green Lacewings (Insecta, Neuroptera, Chrysopidae) from the Early Eocene Okanagan Highlands, Western North America". Journal of Paleontology. 87 (1): 123–146. doi:10.1666/12-052R.1. S2CID 130797848.
77. ^ Archibald, S.B.; Makarkin, V.N. (2015). "A new species of Archaeochrysa Adams (Neuroptera: Chrysopidae) from the early Eocene of Driftwood Canyon, British Columbia, Canada". The Canadian Entomologist. 147 (4): 359–369. doi:10.4039/tce.2014.53. S2CID 86209088.
78. ^ Archibald, S.B.; Makarkin, V.N. (2017). "A new fossil green lacewing (Neuroptera: Chrysopidae) from the early Eocene Driftwood Canyon, Canada". Zootaxa. 4324 (2): 397–400. doi:10.11646/zootaxa.4324.2.12. S2CID 90888385.
79. ^ Archibald, S. B.; Makarkin, V. N. (2015). "The second genus and species of the extinct neuropteroid family Corydasialidae, from early Eocene McAbee, British Columbia, Canada: do they belong to Megaloptera?". Zootaxa. 4040 (5): 569–575. doi:10.11646/zootaxa.4040.5.5. PMID 26624691. S2CID 2034844.
80. ^ a b Makarkin, V. N.; Archibald, S. B.; Oswald, J. D. (2003). "New Early Eocene brown lacewings (Neuroptera: Hemerobiidae) from western North America". The Canadian Entomologist. 135 (5): 637–653. CiteSeerX 10.1.1.489.5852. doi:10.4039/n02-122. S2CID 53479449.
81. ^ Makarkin, V. N.; Wedmann, S.; Weiterschan, T. (2016). "A new genus of Hemerobiidae (Neuroptera) from Baltic amber, with a critical review of the Cenozoic Megalomus-like taxa and remarks on the wing venation variability of the family". Zootaxa. 4179 (3): 345–370. doi:10.11646/zootaxa.4179.3.2. PMID 27811679.
82. ^ Makarkin, V. N. (2023). "Fossil Hemerobiidae (Neuroptera) from the Eocene Tadushi Formation, the Russian Far East, with description of a new genus". Zootaxa. 5297 (1): 115–123. doi:10.11646/zootaxa.5297.1.6.
83. ^ Makarkin, V. N.; Archibald, S. B. (2009). "A new genus and first Cenozoic fossil record of moth lacewings (Neuroptera: Ithonidae) from the Early Eocene of North America" (PDF). Zootaxa. 2063: 55–63. doi:10.11646/zootaxa.2063.1.3. S2CID 13922025.
84. ^ a b c d e f g h Archibald, S.B.; Makarkin, V.N. (2006). "Tertiary Giant Lacewings (Neuroptera: Polystechotidae): Revision and Description of New Taxa From Western North America and Denmark". Journal of Systematic Palaeontology. 4 (2): 119–155. doi:10.1017/S1477201906001817. S2CID 55970660.
85. ^ Meunier, F. (1897). "Observations sur quelques insectes du Corallien de la Bavière". Rivista Italiana di Paleontologia. 3: 18–23.
86. ^ Shcherbakov, D. E. (2006). "The earliest find of Tropiduchidae (Homoptera: Auchenorrhyncha), representing a new tribe, from the Eocene of Green River, USA, with notes on the fossil record of higher Fulgoroidea". Russian Entomological Journal. 15: 315–322.
87. ^ Scudder, S. H (1895). "Canadian fossil insects, myriapods and arachnids, 1. The Tertiary Hemiptera of British Columbia". Geological Survey of Canada Contributions to Canadian Palaeontology. 2: 5–26.
88. ^ Archibald, S.B.; Makarkin, V.N. (2020). "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". The Canadian Entomologist. 152 (3): 269–287. doi:10.4039/tce.2020.10. S2CID 216238397.
89. ^ Archibald, S. B.; Makarkin, V. N.; Ansorge, J. (2009). "New fossil species of Nymphidae (Neuroptera) from the Eocene of North America and Europe" (PDF). Zootaxa. 2157: 59–68. doi:10.11646/zootaxa.2157.1.4. S2CID 9304328.
90. ^ a b c Makarkin, V.N.; Archibald, S.B.; Mathewes, R.W. (2021). "New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene of western North America, with taxonomic remarks". Zootaxa. 4980 (1): 142–156. doi:10.11646/zootaxa.4980.1.9. PMID 34186986. S2CID 235685548.
91. ^ Makarkin, V.; Archibald, S.B. (2014). "An unusual new fossil genus probably belonging to the Psychopsidae (Neuroptera) from the Eocene Okanagan Highlands, western North America". Zootaxa. 3838 (3): 385–391. CiteSeerX 10.1.1.692.1185. doi:10.11646/zootaxa.3838.3.8. PMID 25081783.
92. ^ a b c d e f Archibald, S. B.; Cannings, R. A. (2019). "Fossil dragonflies (Odonata: Anisoptera) from the early Eocene Okanagan Highlands, western North America". The Canadian Entomologist. 151 (6): 783–816. doi:10.4039/tce.2019.61.
93. ^ a b c d e f g h i j k l m n o 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". Zootaxa. 4934 (1): zootaxa.4934.1.1. doi:10.11646/zootaxa.4934.1.1. PMID 33756770.
94. ^ Archibald, S. B.; Cannings, R. A. (2021). "A new genus and species of Euphaeidae (Odonata, Zygoptera) from the early Eocene Okanagan Highlands locality at Republic, Washington, U.S.A.". Zootaxa. 4966 (3): 392–400. doi:10.11646/zootaxa.4966.3.11. PMID 34186607. S2CID 235557114.
95. ^ a b Archibald, S. B.; Gu, J.-J.; Mathewes, R. W. (2022). "The Palaeorehniidae (Orthoptera, Ensifera, "Zeuneropterinae"), and new taxa from the early Eocene Okanagan Highlands, western North America". Zootaxa. 5100 (4): 559–572. doi:10.11646/zootaxa.5100.4.6. PMID 35391059. S2CID 247046978.
96. ^ 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". The Canadian Entomologist. 147 (6): 744–753. doi:10.4039/tce.2015.2. S2CID 86608533.
97. ^ Vladimir N. Makarkin; S. Bruce Archibald; James E. Jepson (2019). "The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America". The Canadian Entomologist. 151 (4): 521–530. doi:10.4039/tce.2019.26. S2CID 196660529.
98. ^ a b c d e Archibald, S. B.; Makarkin, V. N. (2021). "Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation". Zootaxa. 4951 (1): 41–79. doi:10.11646/zootaxa.4951.1.2. PMID 33903413. S2CID 233411745.
99. ^ Hussakof, L. (1916). "A new cyprinid fish, Leuciscus rosei, from the Miocene of British Columbia". American Journal of Science. 4 (247): 18–20. Bibcode:1916AmJS...42...18H. doi:10.2475/ajs.s4-42.247.18.
100. ^ a b c d e Wilson, M. V. (1978). "Eohiodon woodruffi n. sp.(Teleostei, Hiodontidae), from the Middle Eocene Klondike Mountain Formation near Republic, Washington". Canadian Journal of Earth Sciences. 15 (5): 679–686. Bibcode:1978CaJES..15..679W. doi:10.1139/e78-075.
101. ^ a b Wilson, M.V.H. (1979). "A second species of Libotonius (Pisces: Percopsidae) from the Eocene of Washington State". Copeia. 1979 (3): 400–405. doi:10.2307/1443214. JSTOR 1443214.
102. ^ a b c d e f Wilson, MVH (1982). "A new species of the fish Amia from the Middle Eocene of British Columbia". Palaeontology. 25 (2): 413–424.
103. ^ a b c Grande, L.; Bemis, W. E. (1998). "A comprehensive phylogenetic study of amiid fishes (Amiidae) based on comparative skeletal anatomy. An empirical search for interconnected patterns of natural history". Journal of Vertebrate Paleontology. 18 (sup1): 1–696. doi:10.1080/02724634.1998.10011114.
104. ^ a b c d Liu, J. (2021). "Redescription of 'Amyzon' brevipinne and remarks on North American Eocene catostomids (Cypriniformes: Catostomidae)". Journal of Systematic Palaeontology. 19 (9): 677–689. doi:10.1080/14772019.2021.1968966. S2CID 238241095.
105. ^ a b Bruner, JC (1991). "Comments on the Genus Amyzon (Family Catostomidae)". Journal of Paleontology. 65 (4): 678–686. doi:10.1017/S0022336000030766. S2CID 127768517.
106. ^ Wilson, M. V. (1996). "Fishes from Eocene lakes of the interior". In R. Ludvigsen (ed.). Life in stone: a natural history of British Columbia's fossils. Vancouver, BC: The University of British Columbia Press. pp. 212–224.
107. ^ a b c d Hilton, E.; Grande, L. (2008). "Fossil Mooneyes (Teleostei: Hiodontiformes, Hiodontidae) from the Eocene of western North America, with a reassessment of their taxonomy". Geological Society, London, Special Publications. 295 (1): 221–251. Bibcode:2008GSLSP.295..221H. doi:10.1144/SP295.13. S2CID 128680041.
108. ^ Wilson, M.V.H.; Li, Guo-Qing (1999). "Osteology and systematic position of the Eocene salmonid †Eosalmo driftwoodensis Wilson from western North America" (PDF). Zoological Journal of the Linnean Society. 99 (125): 279–311. doi:10.1111/j.1096-3642.1999.tb00594.x. Retrieved 2010-01-01.
109. ^ a b Stockey, R.; Pigg, K. (1994). "Vegetative growth of Eorhiza arnoldii Robison and Person from the Middle Eocene Princeton chert locality of British Columbia". International Journal of Plant Sciences. 155 (5): 606–616. doi:10.1086/297199. S2CID 85094707.
110. ^ Mathewes, R. W.; Greenwood, D. R.; Archibald, S. B. (2016). "Paleoenvironment of the Quilchena flora, British Columbia, during the Early Eocene Climatic Optimum" (PDF). Canadian Journal of Earth Sciences. 53 (6): 574–590. Bibcode:2016CaJES..53..574M. doi:10.1139/cjes-2015-0163. hdl:1807/71979.
111. ^ LePage, B. A.; Currah, R. S.; Stockey, R. A. (1994). "The fossil fungi of the Princeton chert". International Journal of Plant Sciences. 155 (6): 828–836. doi:10.1086/297221. S2CID 85107282.
112. ^ a b c d e f g h i j Mayr, G.; Archibald, S.B.; Kaiser, G.W.; Mathewes, R.W. (2019). "Early Eocene (Ypresian) birds from the Okanagan Highlands, British Columbia (Canada) and Washington State (USA)". Canadian Journal of Earth Sciences. 56 (8): 803–813. Bibcode:2019CaJES..56..803M. doi:10.1139/cjes-2018-0267. S2CID 135271937.
113. ^ a b c d Eberle, J.J.; Rybczynski, N.; Greenwood, D.R. (2014). "Early Eocene mammals from the Driftwood Creek beds, Driftwood Canyon Provincial Park, northern British Columbia". Journal of Vertebrate Paleontology. 34 (4): 739–746. doi:10.1080/02724634.2014.838175. S2CID 130316011.
114. ^ Russell, L.S. (1935). "A middle Eocene mammal from British Columbia". American Journal of Science. 29 (169): 54–55. Bibcode:1935AmJS...29...54R. doi:10.2475/ajs.s5-29.169.54.
115. ^ Dunlop, J.A.; Braddy, S.J. (2011). "Cteniza bavincourti and the nomenclature of arachnid-related trace fossils". The Journal of Arachnology. 39 (2): 250–257. doi:10.1636/CHa10-70.1. S2CID 55206472.
116. ^ a b c d e Scudder, S. H (1878). "Additions to the Insect-fauna of the Tertiary beds at Quesnel, British Columbia". Geological Survey of Canada, Report of Progress for. 1876–1877: 457–464.
117. ^ a b c d e f g h i j k l m n o p q r s t u Scudder, S. H (1877). "Appendix to Mr. George M. Dawson's report. The insects of the Tertiary beds at Quesnel". Geological Survey of Canada, Report of Progress for. 1875–1876: 266–280.
118. ^ a b Michelsen, V. (1996). "First reliable record of a fossil species of Anthomyiidae (Diptera), with comments on the definition of recent and fossil clades in phylogenetic classification". Biological Journal of the Linnean Society. 58 (4): 441–451. doi:10.1111/j.1095-8312.1996.tb01445.x. S2CID 85913252.
119. ^ Radchenko, A. G.; Perkovsky, E. E. (2016). "The ant Aphaenogaster dlusskyana sp. nov. (Hymenoptera, Formicidae) from the Sakhalin amber—the earliest described species of an extant genus of Myrmicinae". Paleontological Journal. 50 (9): 936–946. doi:10.1134/S0031030116090136. S2CID 90471180.
120. ^ Dalla Torre, K. W. (1893). Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7\. Formicidae (Heterogyna). Leipzig: W. Engelmann. p. 289.
121. ^ McKenzie McAnally, L. (1996). "Paleogene mammals on land and at sea". In Ludvigsen, R. (ed.). Life in Stone: A Natural History of British Columbia's Fossils. UBC Press. pp. 208–209 & fig 16.4. ISBN 978-0774841511.