The Paleobiota of the Klondike Mountain Formation comprises a diverse suite of Early Eocene plants and animals recovered from North Central Washington State. The formation outcrops in locations across the north western area of Ferry County, with major sites in Republic, north west of Curlew Lake, and on the Toroda Creek area. The formation is the southern most of the Eocene Okanagan Highlands, sharing much of the paleoflora and paleofauna with site across Central and southern British Columbia.
Dillhoff et al. (2013) reference undescribed moss specimens known from the Klondike Mountain Formation known from vegetative gametophytes and they noted them to be similar to undescribed specimens from the Allenby Formation and Horsefly shales.[1]
Undescribed
Undescribed[1]
Undescribed moss specimens
Rare specimens of Selaginella fossils were noted by Wehr (1998), with no species level description.[2]
Selaginellaceae
Selaginella
Undescribed[2]
A spikemoss Not described to species
Cystopteridaceae
Cf. Cystopteris
A possible bladder fern relative Not described to species
Dennstaedtiaceae
Dennstaedtia
†Dennstaedtia christophelii[3]
Pigg et al.
A Hayscented fern
Equisetaceae
Equisetum
Undescribed[4][2]
A scouring rush Not described to species
Hymenophyllaceae
Hymenophyllum
†Hymenophyllum axsmithii[3]
A filmy fern
Lygodiaceae
Lygodium
Undescribed[2][5]
A climbing fern Not described to species
Salviniaceae
Azolla
†Azolla primaeva[6]
Arnold
A mosquito fern
Salvinia
Undescribed[7]
A "watermoss" species.Not described to species Found only in Curlew half graben sites.
Three major groups of gymnosperms are present in the Klondike Mountain Formation, with the most speciose being the pinophytes. The ginkgophytes are represented by two species pf Ginkgo, while an undescribed Zamiaceae member is the sole cycadophyte.
Zamiaceae
Undescribed[8][9]
A zamiaceous cycad. Not described to genus/species
Ginkgo biloba[10]
Linnaeus
A ginkgo
†Ginkgo dissecta[10]
Mustoe
Calocedrus
Undescribed[9][11]
An incense cedarNot described to species
Chamaecyparis
Undescribed[12][13]
A false cypressNot described to speciesPossibly in the Callitropsis nootkatensis lineage.[14][13]
Cryptomeria
Undescribed[12][5]
A sugiNot described to species
Glyptostrobus
Undescribed[12]
A Chinese swamp cypressNot described to species
Metasequoia
†Metasequoia occidentalis[15][16]
(Newberry) Chaney
A dawn redwood
Sequoia
†Sequoia affinis[15]
Lesquereux
A coast redwood Reported as "Sequoia langsdorfii" by Brown, 1935
Taiwania
A Taiwania speciesnot described to species
Taxodium
†Taxodium dubium[17][15]
(Sternberg) Heer
A bald cypress
Thuja
An arborvitaeNot described to species
Thujopsis?
Undescribed[13]
A possible ThujopsisNot described to species.
Abies
†Abies milleri[18]
Shorn & Wehr
Oldest true fir described
Picea
A spruce Not described to species
†Pinus latahensis[17]
Berry
A 5-needle pine
†"Pinus macrophylla"[17]
A 3-needle pine, jr homonym to Pinus macrophylla Lindley 1839
†Pinus monticolensis[17]
A pine seed morphogenus
†Pinus tetrafolia[17]
A purported 4-needle pine Noted by Berry as "highly improbable that this should represent a distinct botanic species"
Pseudolarix
†Pseudolarix wehrii[19]
Gooch
A golden larch Originally described as Pseudolarix americana.[19]
Tsuga
A hemlockNot described to species
Sciadopityaceae
Sciadopitys
An umbrella pine Not described to species
Amentotaxus
A yew Not described to species
Cephalotaxus
A yewOriginally placed in the Miocene Cephalotaxus bonseri[17][20] Not described to species
Taxus
A yewNot described
Angiosperms, commonly called flowering plants belong to a single plant clade which diverged from other plants during the prior to the Cretaceous, and began to rapidly evolve and radiate by the Middle Cretaceous.[21] Angiosperm diversification during the Cretaceous and Paleocene resulted in eight recognized clades that are segregated into two groups the Basal angiosperms and Core angiosperms. Present in the Klondike Mountain Formation are four of the eight groups, Nymphaeales representing Basal Angiosperms, plus Magnoliids, Monocots, and Eudicots all in the Core angiosperms.
The Basal Angiosperms are represented by a single Nymphaeales water-lily species Nuphar carlquistii,[22] though a second member, Allenbya collinsonae, has been described from the Princeton Chert.[23] Wehr (1995) illustrated two fossils that were tentatively identified as fruits of the banana genus Ensete and the extinct myrtle genus Paleomyrtinaea respectively,[24] however further fossil finds resulted in the re-identification of the first as a N. carlquistii rhizome section, and the second is a seed mass from the same water-lily.[22]
Nymphaeaceae
Nuphar
†Nuphar carlquistii[22]
DeVore, Taylor, & Pigg
A waterlily, Rhizome sections were first identified as Ensete Seed masses first identified as Paleomyrtinaea.
Under the APG IV system of flowering plant classification, the magnoliids are divided into four orders Canellales, Laurales, Magnoliales, and Piperales. Member species and undescribed taxa placed confidently in the Laurales and Magnoliales are present in the formation. The laurales are the most diverse magnoliid order of the formation with one described species Sassafras hesperia plus three tentatively identified genera which have not been described. Of the magnoliales, only an undescribed Magnolia, having possible affinity with Magnolia subg. Talauma, is found in the formation, while Liriodendroxylon princetonensis has described from permineralized wood in the Princeton Chert.[25] The extinct angiosperm genus Dillhoffia has noted similarities to the piperalean family Aristolochiaceae, but was left incertae sedis as to family by Manchester and Pigg (2008) due to a lack of confident morphological characters for placement. Piperales are known from the Princeton chert, with Saururus tuckerae representing the oldest confident Saururaceae species in the fossil record.[26]
†Litseaphyllum
A lauraceous form species. Not described to species
Ocotea
A stinkwood species. Not described to species
Phoebe
Undescribed[16]
A Phoebe species. Not described to species
Sassafras
†Sassafras hesperia[16]
A sassafras
Magnoliaceae
Magnolia
Undescribed[27]
A magnolia, possibly Magnolia subgenus Talauma Not described to species
incertae sedis
†Dillhoffia
†Dillhoffia cachensis[28]
Manchester & Pigg
A flower of uncertain floral affinity
The second largest clade of flowering plants, monocots are divided into eleven separate orders and of those, the Alismatales, Asparagales, Liliales, and Poales are found in the Klondike Mountain Formation, each represented by a single taxon. The Alismatales are represented by the Araceae species Orontium wolfei, which is considered similar to the modern golden clubs of eastern North America, while the extinct Paleoallium belongs to the Liliales. Asparagales and Poales are both present as undescribed species associated with the genera Smilax and Typha respectively. Extinct genera of monocots are also represented in the Princeton chert by the arecalean palm Uhlia,[29] the alismatalean genus Heleophyton,[30] the alismatalean Keratosperma,[31] the asparagalean pollen morphogenus Pararisteapollis,[32] the lilialean genus Soleredera,[33] and the poalean genus Ethela,[34]
Amaryllidaceae
†Paleoallium
†Paleoallium billgenseli[35]
Pigg, Bryan, & DeVore
An onion relative
Araceae
Orontium
†Orontium wolfei[36]
Bogner, Johnson, Kvaček & Upchurch
A golden club
Smilacaceae
Smilax
A greenbrier species.Not described to species.
Typhaceae
Typha
A cattail species.Not described to species.
Over a dozen different Rosaceae genera, both extant and extinct, have been identified in the formation providing some of the oldest reliable macrofossil records (excluding fossil pollen) for the family.[37] Benedict et al. (2011) described first fossils for the prunoid genus Oemleria along with the oldest Prunus flowers. The Prunus flowers are complemented by leaf fossils representing five to six distinct morphotypes.[38] Spiraea is known from an inflorescence with multiple flowers and leaves that are either from the genus or a closely related extinct type. The leaves frequently are preserved with a persistent stipule, a feature not found in modern Spiraea species. The firethorn genus Pyracantha and the South American genus Hesperomeles have been tentatively identified from leaves while Maloidea leaves belonging to either Malus or Pyrus have been found. Two distinct species of the Asian endemic genus Photinia are known, however only one of them Photinia pagae had been described as of 2007.[38] The rosaceous genus Physocarpus had been reported by Hopkins and Wehr (1994) as also occurring in the formation,[27] however subsequent examination of the fossils by Oh & Potter (2005) failed to find stellate trichomes which are a distinct feature of the genus. They noted the fossils might be stem Neillieae, the rose tribe containing both Physocarpus and Neillia, or possibly Rubus, Crataegus, or Ribes.[39]
Fossils of both Sorbus and Rhus species leaves showing evidence of being interspecies hybrids have been noted from the formation and Flynn, DeVore and Pigg (2019) described four species of sumac which formed multiple hybrids.[40] Between three and four Trochodendraceae species that have been described from the Klondike Mountain Formation. Broadly circumscribed four species in three genera have been identified at Republic, Paraconcavistylon wehrii, Pentacentron sternhartae, Tetracentron hopkinsii, and Trochodendron nastae. Additionally the species Trochodendron drachukii is known from related Kamloops group shales at the McAbee Fossil Beds near Cache Creek, British Columbia. Manchester et al. 2018 noted that Tr. drachukii is likely the fruits of Tr. nastae, while Pe. sternhartae are likely the fruits of Te. hopkinsii.[41] If fossils of the fruits and foliage in attachment are found, that would bring the species count down to three whole plant taxa.[41] Additionally, the extinct genus Nordenskioldia is also known from the formation. The placement of Nordenskioldia is debated, with some treatments placing it into Trochodendraceae, while a 2020 analysis placed it outside of the crown-group Trochodendaceae.[42] Wesley Wehr in 1994 reported Bignoniaceae seeds along with a single Rubiaceae fruit and an isolated Fabaceae leaf.[43] An update of the floral list by Wehr and Manchester published in 1996 added an additional fifteen taxa identified from reproductive structures such as flowers fruits or seeds.[24]
Pigg, Manchester, and Wehr (2003) noted in during the description of Corylus johnsonii and Carpinus perryae that they were the oldest confirmed hazelnut and hornbeam fossils.[44] That status was affirmed by Forest et al. (2005) who used both as fossil calibration points for phylogenetic analysis of Betulaceae.[45] Within the family Bignoniaceae, the fossil seeds and fruits are noted by Ze-Long Nie et al (2006) as the oldest confirmed for the family.[46]
†Rhus boothillensis[40]
Flynn, DeVore, & Pigg
A sumac, Hybridized with other Klondike Mountain Formation Rhus
†Rhus garwellii[40]
†Rhus malloryi[16][40]
(Wolfe and Wehr) Flynn, DeVore & Pigg
†Rhus republicensis[40]
Aquifoliaceae
Ilex
Undescribed[5]
A holly Not described to species
Araliaceae
Aralia
Undescribed[9]
A spikenard speciesNot described to species.
Alnus
†Alnus parvifolia[16]
(Berry) Wolfe & Wehr
An Alder
Betula
†Betula leopoldae[16][47]
Wolfe & Wehr
A birch
Carpinus
†Carpinus perryae[44]
Pigg, Manchester, & Wehr
A hornbeam
Corylus
†Corylus johnsonii[44]
A hazel nut
†Palaeocarpinus barksdaleae[44]
A birch relative
Undescribed[44]
A birch relative different from the other Okanagan Highlands species
Bignoniaceae
Undescribed[43]
A catalpa family member Not described to species
Burseraceae
†Barghoornia
†Barghoornia oblongifolia[16]
An extinct Bursera relative
Celtidaceae
Pteroceltis
Undescribed[48]
A cannabaceous fruitNot described to species
Cercidiphyllaceae
Cercidiphyllum
†Cercidiphyllum obtritum[16]
(Dawson) Wolfe & Wehr
A katsura with suggested affinity to †Joffrea,[9] First described as "Populus" obtrita
Cornaceae
Cornus
Undescribed[16][9][5]
A dogwood species,Not described to species
Elaeocarpaceae
Sloanea
An elaeocarpaceous fruitNot described to species
Arbutus
Undescribed[27][5]
A madrone relativeNot described to species
Rhododendron
Undescribed[27][9][5]
A rhododendronNot described to species
Cf. Leucothoe
A doghobble relativeNot described to species
Eucommiaceae
Eucommia
†Eucommia montana[49][5]
Brown
A "hard rubber tree"
†Castaneophyllum
A Castanea relativeNot described to species
†Fagopsis
†Fagopsis undulata[16]
(Knowlton) Wolfe & Wehr
A beech relative
Fagus
†Fagus langevinii[50]
Manchester & Dillhoff
A beech
Quercus
Undescribed[43][9][5]
An oakNot described to species
"Species 1"[27][9][5]
A currentNot described to species
"Species2"[27]
Corylopsis
†Corylopsis reedae[27][51]
Radtke, Pigg, & Wehr
A winter-hazel
Fothergilla
†Fothergilla malloryi[51]
A witch alder
Hydrangea
A Hydrangea. Not described to species
Philadelphus
A mock-orangeNot described to species
Icacinaceae
†Palaeophytocrene
Unidentified[43][9]
A Phytocrene relativeNot described to species
Iteaceae
Itea
Undescribed[16][5]
A virginia willow speciesNot described to species.
Carya
Undescribed[9][5]
A walnut family relative.Not described to species.
†Cruciptera
†Cruciptera simsonii[52]
(Brown) Manchester
A walnut family relative.
Juglans
Pterocarya
A wing nutNot described to species
Lythraceae
Decodon
A swamp loosestrife Not described to species
Craigia
A Craigia speciesNot described to species
†Florissantia
†Florissantia quilchenensis[53]
(Mathewes & Brooke) Manchester
A chocolate relative
Hibiscus
A hibiscusNot described to species
†Plafkeria
A linden relativeNot described to species
Tilia
†Tilia johnsoni[16]
A Linden
Cf. Tilia
Linden relative fruitsNot described to species
Melastomataceae?
"Schoepfia"
†"Schoepfia" republicensis[16]
(LaMotte) Wolfe & Wehr
First described as a dogwood under "Cornus acuminata then a possible Schoepfia species,Placement in Schoepfiaceae rejected by Malécot and Lobreau‐Callen, (2005)[54]S. republicensis fossil figured as Melastomataceae by Renner et al (2001)[55]
Menispermaceae
Calycocarpum
Undescribed[24]
A moonseedNot described to species
Moraceae
Morus
A mulberry, two types known. Not described to species
Myricaceae
Comptonia
†Comptonia columbiana[16]
Dawson
A Comptonia
Nyssaceae
†Tsukada
†Tsukada davidiifolia[16]
A dove-tree relative
Oxalidaceae?
†Averrhoites
A leaf morphotype of uncertain affiliation.First described as visually similar to AverrhoaNot described to species.
†Langeranthus
†Langeranthus dillhoffiorum[56]
Huegele & Manchester
A plane tree fruit taxon affiliated with Langeria
†Langeria
†Langeria magnifica[16]
A plane treeFormerly identified as a witch hazel relative
†Macginicarpa
Manchester
A plane tree fruit taxonNot described to species
†Macginitiea
†Macginitiea gracilis[16]
(Lesquereux) Wolfe & Wehr
A plane tree relative
†Platananthus
A sycamore stamen head isolated stamen clusters placed as Macginistemon Not described to species
Polygalaceae
†Deviacer
A milkwort relativeNot described to species
Ranunculaceae
Clematis
A Clematis Not described to species
Amelanchier
A service berry Not described to species
Cf. Crataegus
A hawthorn relativeNot described to species
Malus
An appleNot described to species
Cf. Malus
Undescribed[38]
A maloid species possibly apple or pear Not described to species
Neviusia
A snow-wreathNot described to species
Oemleria
†Oemleria janhartfordae[37]
Benedict, DeVore, & Pigg
An Osoberry
Photinia
†Photinia pageae[16]
A Christmas-berry relative
Aff. Physocarpus
A possible nine-bark Not described to species Possibly stem Neillieae[39]
†Prunus cathybrownae[37]
A cherry relative
"Species 1"[16]
A prunoid leafNot described to species
"Species 2"[16]
"Species 3"[16]
Pyracantha
A firethorn sp.[38]Tentative record, Not described to species.
Cf. Pyrus
Rubus
A blackberryNot described to species
Aff. Sorbus
A rowan relativeNot described to species.
Spiraea
A bridal wreath Not described to species.
Cf. Spiraea
A bridal wreath relative Not described to species
Meliosma
A Meliosma speciesNot described to species
Sabia
A Sabia speciesNot described to species
Populus
A cottonwoodNot described to species First identified as †Populus lindgreni[17]
†Pseudosalix
A willow relativeNot described to species
Salix
A willowNot described to species
Acer?
†"Acer" arcticum[57]
Heer, 1876
A possible maple
†Acer hillsi[57]
Wolfe & Tanai
A maple
†Acer republicense[57]
†Acer spitzi[57]
†Acer stonebergae[57]
†Acer toradense[57]
†Acer washingtonense[57]
†Acer wehri[57]
Aesculus
Undescribed[9][58]
A horse chestnut Not described to species.
Cf. Boniodendron
†"Koelreuteria" arnoldii[16][59]
Becker
A sapindaceous speciesfirst described as a Koelreuteria species, considered Cf. Boniodendron by Wang et al. (2012).
†Bohlenia
†Bohlenia americana[16]
(Brown) Wolf & Wehr
An extinct sapindalean species
Dipteronia
†Dipteronia brownii[60]
McClain & Manchester
A Dipteronia
Koelreuteria
†Koelreuteria dilcheri[59]
Wang et al.
A Koelreuteria species
Schisandraceae
Kadsura
A kadsura species. Not described to species
†Ternstroemites
"Species A"[16]
A theaceous species similar to GordoniaNot described to species
"Species B"[16]
A theaceous species similar to CleyeraNot described to species
Trochodendraceae?
†Nordenskioldia
A trochodendroid of uncertain placement.[42]Fruits of the leaf taxon ZizyphoidesNot described to species.
†Paraconcavistylon
†Paraconcavistylon wehrii[41][42]
(Manchester et al.)
A Trochodendralefirst described as "Concavistylon" wehriimoved to a new genus in 2020.
†Pentacentron
†Pentacentron sternhartae[41]
Manchester et al.
A Trochodendrale
Tetracentron
†Tetracentron hopkinsii[41]
A Trochodendrale,possibly the leaves of Pentacentron sternhartae
Trochodendron
†Trochodendron nastae[61]
Pigg, Wehr, & Ickert-Bond
A TrochodendronPossibly the leaves of Trochodendron drachukii
†Zizyphoides
A trochodendroid of uncertain placement.[42]Leaves of the fruit taxon NordenskioldiaNot described to species.
†Cedrelospermum
An elm relativeNot described to species
†Ulmus chuchuanus[62]
(Berry) LaMotte
An elm species Leaves with features of Ulmus subg. Ulmusfruits with features of Ulmus subg. Oreoptelea
†Ulmus okanaganensis
Denk & Dillhoff
An elm species, the fruits were first identified as Ulmus section Chaetoptelea.
Vitaceae
Vitis
Undescribed[27][43]
Grape seeds[43] and leaves[27]Not described to species
†Calycites
†Calycites ardtunensis[43]
Crane
A winged fruit of unidentified affinities
†Pteroheterochrosperma
†Pteroheterochrosperma horseflyensis[63]
Smith, Greenwalt & Manchester
A samara of uncertain affiliation.
†Pteronepelys
†Pteronepelys wehrii[64]
A samara of uncertain affinities.
†Republica
†Republica hickeyi[16]
An incertae sedis angiosperm possibly of Hamamelididae affiliations
The insect fauna of the Klondike Mountain Formation includes representatives from over 13 orders, based on a 1992 estimate, including immature though adult specimens and both terrestrial and aquatic taxa.[65] The most prevalent orders are Diptera and Hemiptera, each making up approximately 30% of the fossil insects known in 1992.
Blaberidae
A Diplopterine cockroach Not described to genus/species
Blattoidae
Undescribed[65]
A blattoidean cockroachNot described to genus/species
Isoptera
Undescribed termites of uncertain affiliation
A list of Coleopteran families identified by 1992 included Carabidae, Cerambycidae, Chrysomelidae, Curculionidae, Dytiscidae, Elateridae and Lucanidae,[65] but the beetle fauna has not been described in depth yet, with only two weevil species having been fully described.[66][67] A third beetle group belonging to the bean beetle tribe Pachymerini has been identified as palm beetles of the Caryobruchus–Speciomerus genus group.[68]
†Eoceneithycerus
†Eoceneithycerus carpenteri[66]
Legalov, 2013
An Ithycerinae weevil
†Ithyceroides
†Ithyceroides klondikensis[67]
Legalov, 2015
Carabidae
A ground beetleNot described to genus/species
Cerambycidae
A long-horn beetleNot described to genus/species
Chrysomelidae
Caryobruchus–Speciomerus genus group
Undescribed[68]
palm beetles in the tribe Pachymerini. Not described to genus/species
Dytiscidae
A diving beetleNot described to genus/species
Elateridae
A click beetleNot described to genus/species
Lucanidae
A stag beetleNot described to genus/species
Unidentified
Undescribed[69]
A possible staphylinoid beetle not described
The order Dermaptera was first reported in 1992[65] and is known from a series of isolated partial specimens, mostly abdominal sections with the distinct anal forceps attached. Based on the forceps structuring the specimens were tentatively assigned to the modern family Forficulidae, as the oldest North American representatives of the family known at that time.[70]
"Forficulid species 1"[70]
A possible forficulid earwig with long cerci Not described to genus/species
"Forficulid species 2"[70]
A possible forficulid earwig with short cerci Not described to genus/species
Bibionidae
A march flyNot described to genus/species
Cecidomyiidae
Undescribed[71]
Trace fossilsCecidomyiid midge galling on various host leavesNot described to genus/species
Empididae
A dagger flyNot described to genus/species
Mycetophilidae
A fungus gnatNot described to genus/species
Pipunculidae
†Metanephrocerus
†Metanephrocerus belgardeae[72]
Archibald, Kehlmaier, & Mathewes, 2014
A pipunculid big-headed fly
Syrphidae
Unidentified[73]
A hover flyNot described to genus/species
Tipulidae
A crane flyNot described to genus/species
Lewis (1992) listed one species of Heptageniidae and three specimens that he did not place to family.[65] 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.[74] 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.[75]
Heptageniidae
Indeterminate
Indeterminate[65]
A flat headed mayfly nymph.Tentatively suggested as Heptagenia or Stenonema by Lewis & Wehr (1993) Deemed indeterminate below family level by Sinitchenkova (1999)[75]
Neoephemeridae
Neoephemera
†Neoephemera antiqua[75]
Sinitchenkova, 1999
A squaregill mayfly
Aphididae
An aphidNot described to genus/species.
Aphrophora
An aphrophorid spittlebugNot described to species.
†Petrolystra
†Palecphora
A cercopid froghopperNot described to species.
A cercopid froghopperNot described to genus/species.
Fulgoroidea
A frog hopperNot described to genus/species.
Pentatomidae
A Shield or stink bugNot described to genus/species
A review of the Okanagan highlands hymenoptera published in 2018 identified four "Symphyta" families in the formation Cimbicidae, Pamphiliidae, Siricidae, and Tenthredinidae. Of the "Apocrita" families thirteen are represented, the "Parasitica" families are Braconidae, Diapriidae Ichneumonidae, Proctotrupidae, and Roproniidae while the Vespoidea families are Formicidae, Pompilidae, Scoliidae and Vespidae. Within Apoidea the "Spheciformes" families include Angarosphecidae and Sphecidae while Halictidae is the sole "Apiformes" family known from body fossils. Prunus and Ulmus leaves have been found having damage that is consistent with the damage pattern left by Megachilidae species bees when they remove sections of tissue for nest lining. There are several additional Apoidea fossils that were left as incertae sedis in the group based on the similarity between them and Paleorhopalosoma menatensis, a Paleocene species described from the Menat Formation Auvergne, France. The placement of P. menatensis is uncertain, having been initially described as a member of Rhopalosomatidae, but is possibly an Angarosphecidae or closely related taxon, based on the wing and body morphology.[76]
†Eosphecium
Undescribed[76]
An angarosphecid spheciform wasp. Not described to species
An angarosphecid spheciform wasp Likely not Eosphecium. Not described to species
Braconidae
Undescribed[65][76]
braconid parasitic waspsNot described to genus/species.
†Leptostigma alaemacula[76][77]
Archibald & Rasnitsyn, 2023
A cenocimbicine cimbicid sawfly.
†Leptostigma brevilatum[76][77]
A cenocimbicine cimbicid sawfly Tentatively identified from Republicdescribed from the McAbee fossil beds
Cynipidae
Trace fossilscynipid Cynipoid gallwasp galling on various host leavesNot described to genus/species
Diapriidae
A diapriid diaprioid wasp Not described to species
†Klondikia
†Klondikia whiteae[78]
Dlussky & Rasnitsyn, 2003
An ant of uncertain subfamily affiliation
Myrmeciites
"Indesterminate"[79]
Archibald, Cover, & Moreau, 2006
A bulldog ant form genus
Oecophylla
†Oecophylla kraussei[80]
(Dlussky & Rasnitsyn, 1999)
An ant, described as Camponotites kraussei, Moved to Oecophylla kraussei in 2017[81]
†Propalosoma
†Propalosoma gutierrezae[80]
Dlussky & Rasnitsyn, 1999
A bulldog ant, first described as a Rhopalosomatidae wasp, moved to myrmeciinae in 2018[82]
Ants of uncertain subfamily placement.[76]
Halictidae (?)
A possible sweat bee Not described to genus/species
Ichneumonidae
ichneumonid parasitic wasps unplaced to subfamily Not described to genus/species
Megachilidae
Megachilid leaf-cutter bee herbivory trace fossils on leaves Not described to genus/species
Pamphiliidae
†Ulteramus
†Ulteramus republicensis[83]
Archibald & Rasnitsyn, 2015
A parasitic wasp
Pompilidae
A pompilid spider wasp Not described to genus/species
Proctotrupidae
A proctotrupid parasitic wasp Not described to genus/species
Roproniidae
A roproniid (sensu lato) proctotrupoid wasp Not described to genus/species
Siricidae
†Eourocerus
†Eourocerus anguliterreus[84]
Archibald & Rasnitsyn, 2022
A siricine horntail.
Scoliidae
An archaeoscoliine scoliid wasps Not described to genus/species
Sphecidae
A sphecid (sensu stricto) wasp Not described to genus/species
Tenthredinidae
A Tenthredinid sawfly Not described to genus/species
Vespidae
A vespid wasp Not described to genus/species
A solitary lepidopteran body fossil has been recovered, but no full descriptive work has been made on the specimen, aside from a single PhD dissertation. 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.[85] Trace fossil evidence from leaf fossil herbivory indicates at least four other possible lepidopteran families were present in the formation. The
Coleophoridae
Trace fossilsColeophorid hole feeding and larval casesNot described to genus/species
Erebidae
Undescribed[85]
An arctiine tiger moth Not described
Heliozelidae
Undescribed[71][86]
Trace fossilsheliozelid leaf mining similar to Antispila minesNot described to genus/species
Incurvariidae
Aff. Incurvaria
Trace fossilsincurvariid leaf mining similar to IncurvariaNot described to genus/species
Nepticulidae
Stigmella
Trace fossilsnepticulid leaf mining referred to StigmellaNot described to genus/species
A number of mecopteran species belonging to the families Cimbrophlebiidae, Dinopanorpidae, Eorpidae, and Panorpidae are also known.[87]
†Cimbrophlebia brooksi[87]
Archibald, 2009
A Cimbrophlebiid scorpionfly
†Cimbrophlebia westae[87]
†Dinokanaga andersoni[88]
Archibald, 2005
A scorpion fly species
†Dinokanaga dowsonae[88]
†Dinokanaga sternbergi[88]
†Eorpa elverumi[89]
Archibald, Mathewes, & Greenwood, 2013
A mecopteran scorpionfly
?†Eorpa ypsipeda[89]
A mecopteran scorpionfly, tentatively identified
Panorpidae
Undescribed[89]
Undescribed common scorpionfliesNot described to genus/species
The neuropteran insects (lacewings and their allies) identified as of 2014 include species from the families Berothidae, Chrysopidae, Hemerobiidae, Ithonidae (including Polystoechotidae), Nymphidae, Osmylidae, and Psychopsidae.[90]
Chrysopidae
†Adamsochrysa
†Adamsochrysa wilsoni[91]
Makarkin & Archibald, 2013
2013
A nothochrysine green lacewing
Hemerobiidae
†Archibaldia
†Archibaldia wehri[92][93][94]
(Makarkin, Archibald, & Oswald, 2003)
2003
A hemerobiid lacewing originally placed in †Cretomerobius Moved to †Proneuronema (2016) Moved to †Archibaldia (2023)
†Allorapisma
†Allorapisma chuorum[95]
Makarkin & Archibald, 2009
2009
A moth lacewing
†Palaeopsychops marringerae[96]
Archibald & Makarkin, 2006
2006
A polystechotid group[97] moth lacewing
†Palaeopsychops timmi[96]
†Polystoechotites barksdalae[96]
†Polystoechotites falcatus[96]
†Polystoechotites lewisi[96]
Nymphidae
?Nymphes
†Nymphes? georgei[98]
Archibald, Makarkin, & Ansorge, 2009
A nymphid lacewing, possibly a species of †Epinesydrion[99]
Osmylidae
†Osmylidia
†Osmylidia glastrai[100]
Makarkin, Archibald, & Mathewes, 2021
2021
A protosmyline osmylid lacewing
Psychopsidae?
†Ainigmapsychops
†Ainigmapsychops inexspectatus[90]
Makarkin & Archibald, 2014
2014
A possible psychopsid lacewing
†Antiquiala
†Antiquiala snyderae[101]
Archibald & Cannings, 2019
2019
A darner dragonfly
†Idemlinea
†Idemlinea versatilis[101]
†Ypshna
†Ypshna brownleei[101]
†Dysagrion
†Dysagrion pruettae[102]
Archibald & Cannings, 2021
A Dysagrionine cephalozygopteran odonate
†Dysagrionites
†Dysagrionites delinei[102]
A dysagrionine cephalozygopteran odonate
†Okanagrion dorrellae[102]
†Okanagrion hobani[102]
†Okanagrion liquetoalatum[102]
†Okanagrion threadgillae[102]
†Okanagrion worleyae[102]
†Okanopteryx
†Okanopteryx jeppesenorum[102]
†Stenodiafanus
†Stenodiafanus westersidei[102]
Euphaeidae
†Republica weatbrooki[103]
A gossamerwing damselfly. Not to be confused with the plant Republica, also from the formation
†Whetwhetaksidae
†Whetwhetaksa
†Whetwhetaksa millerae[102]
Archibald & Cannings
A cephalozygopteran odonate
Palaeorehniidae
†Republicopteron
†Republicopteron douseae[104]
Archibald, Gu, & Mathewes
2022
A grasshopper/hump-back grig relative
Fossil wings first described in 2015 were identified as being from Susumanioidea stick-insects, a group that had previously been known from the Jurassic to the Paleocene only.[105] Archibald and Bradler (2015) did not place Eoprephasma into Susumaniidae family, maintaining that known characters of the describe specimens did not match taxa in the family, they instead kept the genus as Susumanioidea incertae sedis. Phylogenetic analysis of Susumanioidea published by Yang et al. (2021) resulted in placement of Eoprephasma as the sister group to Renphasma deep within the Susumaniidae subfamily Susumaniinae. The phylogeny produced by Yang et al. indicated a sister group state with the Cretaceous genus Renphasma of China, and placed both as the most derived of the Susumaniinae taxa.[106]
†Susumaniidae
†Eoprephasma
†Eoprephasma hichensi[105]
Archibald & Bradler, 2015
2015
A Susumaniinae stick insect species
Raphidiidae
†Megaraphidia
†Megaraphidia klondika[107]
Archibald & Makarkin, 2021
A raphidiid snakefly
Trichopterans are known mainly from laraval cases and occasional isolated wings.[108]
Phryganeidae
Unidentified[65][108]
giant caddisfliesNot described to genus/species
Limnephilidae
unidentified[65]
northern caddisfliesNot described to genus/species
Five species of fish have been identified from the formation, four of which are known from skeletal elements, while the fifth is only known from isolated scales.[109] Of the five species, two are unique to the formation, Hiodon woodruffi and Libotonius pearsoni were both described by paleoichthyologist Mark V. H. Wilson in 1978 and 1979 respectively. The other three species, "Amia" hesperia, Amyzon aggregatum, and Eosalmo driftwoodensis, were first described from Okanagan Highlands formations in British Columbia and subsequently also identified from Ferry County fossils. The first notation of fish fossils in the Republic area was by Joseph Umpleby in his 1910 visit to the area, who collected fish near the Tom Thumb Mine, and sent them to the National Museum of Natural History. After examining the fossils, Charles R. Eastman listed the specimens as belonging to the extinct species Amyzon brevipinne in his Fossil fishes in the collection of the United States National Museum.[110] Research tapered off until a series of fish were collected in the Toroda Creek Graben northwest of Republic by Robert Carl Pearson during his early 1960's field mapping for the Geologic map of the Bodie Mountain quadrangle, Ferry and Okanogan Counties, Washington. The fossils were tentatively identified by paleoichthyologist David Dunkle in 1962 and 1965 as members of the genera Amyzon, Tricophanes, Erismatopterus and an undefined salmonid.[111][112] Pearson sent almost all of the specimens collected to the Smithsonian, but the fossils were never accessioned into the collections and are now considered lost. He did retain one fossil from the initial collection which was later donated to the USGS collections. The largest single work on the fish of the Okanagan Highlands was published by Mark Wilson in 1977 and covered fossils collected from the known British Columbian Okanagan Highlands fossil sites of the time.[113] While not covering the Washington State fossils, Wilson named two of the species that are currently recognized from the Klondike Mountain Formation Amyzon aggregatum and Eosalmo driftwoodensis. Additionally scales attributed to the genus Amia were discussed and the genus Libotonius was named from fossils in the Allenby Formation.[114][115] In the late 1960s a collection of fish from near the Tom Thumb Mine in Republic was compiled by resident R. Woodward. During the summers of 1976 and 1977 the University of Alberta conducted field collecting in both the Republic and Toroda Creek areas, along with the donation of the Woodward collection, yielded a number of fossil catostomids, along with a single percopsid, a salmonid, a hiodontid, and an Amia scale. The hiodontids were subsequently described as the species Eohiodon woodruffi in 1978 based on differences between the Tom thumb Tuff fossils and those found in British Columbian sites.[112] A year later the percopsid fossils were also described as Libotonius pearsoni, extending the range of the genus south from the Allenby Formation.[114]
Bird fossils are limited to mostly isolated feathers that are preserved in the finer grained strata of the lake bed, though one partial bird skeleton has also been recovered.
Amiidae
Amia
†"Amia" hesperia
Wilson, 1977
1977
A bowfin, known from isolated scales
Catostomidae
†Amyzon
†Amyzon aggregatum
A sucker
Unidentified[116]
(Wilson, 1977)
A sucker, originally identified as Amyzon aggregatum
Salmonidae
†Eosalmo
†Eosalmo driftwoodensis[117]
1999
A Salmon
Hiodontidae
Hiodon
†Hiodon woodruffi[112]
Wilson, 1978
1978
A mooneye, first described as "Eohiodon" woodruffi.[112]
†Libotoniidae
†Libotonius
†Libotonius pearsoni[114]
Wilson, 1979
1979
A sand roller relative.
incertae sedis (Aves)
"Unnamed"[118]
indeterminate feathers and a skeleton
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