Earth:Paleobiota of the Klondike Mountain Formation

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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.

Plants

Bryophytes

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]

Family Genus Species Authors Year Notes Images

Undescribed

Undescribed

Undescribed[1]

Undescribed moss specimens

undescribed bryophyte

Lycophytes

Rare specimens of Selaginella fossils were noted by Wehr (1998), with no species level description.[2]

Family Genus Species Authors Year Notes Images

Selaginellaceae

Selaginella

Undescribed[2]

A spikemoss
Not described to species

Selaginella species

Pteridophytes

Family Genus Species Authors Notes Images

Cystopteridaceae

Cf. Cystopteris

Undescribed[2]

A possible bladder fern relative
Not described to species

Dennstaedtiaceae

Dennstaedtia

Dennstaedtia christophelii[3]

Pigg et al.

A Hayscented fern

File:Dennstaedtia christophelii holotype SRIC SR 13-004-001 A img1.tif

Equisetaceae

Equisetum

Undescribed[4][2]

A scouring rush
Not described to species

Equisetum species
Not described

Hymenophyllaceae

Hymenophyllum

Hymenophyllum axsmithii[3]

Pigg et al.

A filmy fern

Lygodiaceae

Lygodium

Undescribed[2][5]

A climbing fern
Not described to species

Salviniaceae

Azolla

Azolla primaeva[6]

Arnold

A mosquito fern

Azolla primaeva

Salviniaceae

Salvinia

Undescribed[7]

A "watermoss" species.
Not described to species
Found only in Curlew half graben sites.

Salvinia species
Not described

Gymnosperms

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.

Cycadophytes

Family Genus Species Authors Notes Images

Zamiaceae

Undescribed

Undescribed[8][9]

A zamiaceous cycad.
Not described to genus/species

Undescribed zamiaceous leaf

Gingkophytes

Family Genus Species Authors Notes Images
Ginkgoaceae Ginkgo

Ginkgo biloba[10]

Linnaeus

A ginkgo

Ginkgo biloba

Ginkgo dissecta[10]

Mustoe

A ginkgo

Ginkgo dissecta

Pinophytes

Family Genus Species Authors Notes Images
Cupressaceae

Calocedrus

Undescribed[9][11]

An incense cedar
Not described to species

Chamaecyparis

Undescribed[12][13]

A false cypress
Not described to species
Possibly in the Callitropsis nootkatensis lineage.[14][13]

Cryptomeria

Undescribed[12][5]

A sugi
Not described to species

Glyptostrobus

Undescribed[12]

A Chinese swamp cypress
Not 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

Sequoia affinis

Taiwania

Undescribed[7]

A Taiwania species
not described to species

Taxodium

Taxodium dubium[17][15]

(Sternberg) Heer

A bald cypress

Thuja

Undescribed[12][13]

An arborvitae
Not described to species

Thujopsis?

Undescribed[13]

A possible Thujopsis
Not described to species.

Pinaceae

Abies

Abies milleri[18]

Shorn & Wehr

Oldest true fir described

Abies milleri

Picea

Undescribed[12]

A spruce
Not described to species

Pinus

Pinus latahensis[17]

Berry

A 5-needle pine

Pinus latahensis

"Pinus macrophylla"[17]

Berry

A 3-needle pine, jr homonym to Pinus macrophylla Lindley 1839

"Pinus macrophylla"

Pinus monticolensis[17]

Berry

A pine seed morphogenus

Pinus monticolensis

Pinus tetrafolia[17]

Berry

A purported 4-needle pine
Noted by Berry as
"highly improbable that this should represent a distinct botanic species"

"Pinus tetrafolia"

Pseudolarix

Pseudolarix wehrii[19]

Gooch

A golden larch
Originally described as Pseudolarix americana.[19]

Pseudolarix wehrii

Tsuga

Undescribed[12]

A hemlock
Not described to species

Sciadopityaceae

Sciadopitys

Undescribed[12]

An umbrella pine
Not described to species

Sciadopitys species
Taxaceae

Amentotaxus

Undescribed[12][13]

A yew
Not described to species

Amentotaxus sp.

Cephalotaxus

Undescribed[12]

A yew
Originally placed in the Miocene Cephalotaxus bonseri[17][20]
Not described to species

Taxus

Undescribed[12]

A yew
Not described

Flowering plants

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.

Nymphaeales

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]

Family Genus Species Authors Notes Images

Nymphaeaceae

Nuphar

Nuphar carlquistii[22]

DeVore, Taylor, & Pigg

A waterlily,
Rhizome sections were first identified as Ensete
Seed masses first identified as Paleomyrtinaea.

Magnoliids

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]

Family Genus Species Authors Notes Images
Lauraceae

Litseaphyllum

Undescribed[2]

A lauraceous form species.
Not described to species

Ocotea

Undescribed[2]

A stinkwood species.
Not described to species

Phoebe

Undescribed[16]

A Phoebe species.
Not described to species

Sassafras

Sassafras hesperia[16]

Berry

A sassafras

Sassafras hesperia

Magnoliaceae

Magnolia

Undescribed[27]

A magnolia, possibly Magnolia subgenus Talauma
Not described to species

Magnolia

incertae sedis

Dillhoffia

Dillhoffia cachensis[28]

Manchester & Pigg

A flower of uncertain floral affinity

Dillhoffia cachensis

Monocots

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]

Family Genus Species Authors Notes Images

Amaryllidaceae

Paleoallium

Paleoallium billgenseli[35]

Pigg, Bryan, & DeVore

An onion relative

Paleoallium billgenseli

Araceae

Orontium

Orontium wolfei[36]

Bogner, Johnson, Kvaček & Upchurch

A golden club

Orontium wolfei

Smilacaceae

Smilax

Undescribed[2]

A greenbrier species.
Not described to species.

Typhaceae

Typha

Undescribed[2]

A cattail species.
Not described to species.

Eudicots

Rhus hybrid leaf with lobed terminal leaflet

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 on 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]

Family Genus Species Authors Notes Images
Anacardiaceae Rhus

Rhus boothillensis[40]

Flynn, DeVore, & Pigg

A sumac,
Hybridized with other Klondike Mountain Formation Rhus

Rhus boothillensis

Rhus garwellii[40]

Flynn, DeVore, & Pigg

A sumac,
Hybridized with other Klondike Mountain Formation Rhus

Rhus garwellii

Rhus malloryi[16][40]

(Wolfe and Wehr) Flynn, DeVore & Pigg

A sumac,
Hybridized with other Klondike Mountain Formation Rhus

Rhus malloryi

Rhus republicensis[40]

Flynn, DeVore, & Pigg

A sumac,
Hybridized with other Klondike Mountain Formation Rhus

Aquifoliaceae

Ilex

Undescribed[5]

A holly
Not described to species

Araliaceae

Aralia

Undescribed[9]

A spikenard species
Not described to species.

Betulaceae

Alnus

Alnus parvifolia[16]

(Berry) Wolfe & Wehr

An Alder

Alnus parvifolia

Betula

Betula leopoldae[16][47]

Wolfe & Wehr

A birch

Betula leopoldae

Carpinus

Carpinus perryae[44]

Pigg, Manchester, & Wehr

A hornbeam

Carpinus parryae

Corylus

Corylus johnsonii[44]

Pigg, Manchester, & Wehr

A hazel nut

Corylus johnsonii
Palaeocarpinus

Palaeocarpinus barksdaleae[44]

Pigg, Manchester, & Wehr

A birch relative

Palaeocarpinus barksdaleae

Undescribed[44]

A birch relative
different from the other Okanagan Highlands species

undescribed Palaeocarpinus species

Bignoniaceae

Undescribed

Undescribed[43]

A catalpa family member
Not described to species

Bignoniaceae sp. fruit

Burseraceae

Barghoornia

Barghoornia oblongifolia[16]

Wolfe & Wehr

An extinct Bursera relative

Barghoornia oblongifolia

Celtidaceae

Pteroceltis

Undescribed[48]

A cannabaceous fruit
Not described to species

Pteroceltis species fruit

Cercidiphyllaceae

Cercidiphyllum

Cercidiphyllum obtritum[16]

(Dawson) Wolfe & Wehr

A katsura with suggested affinity to †Joffrea,[9]
First described as "Populus" obtrita

Cercidiphyllum obtritum

Cornaceae

Cornus

Undescribed[16][9][5]

A dogwood species,
Not described to species

Elaeocarpaceae

Sloanea

Undescribed[48]

An elaeocarpaceous fruit
Not described to species

Sloanea sp.
Ericaceae

Arbutus

Undescribed[27][5]

A madrone relative
Not described to species

Rhododendron

Undescribed[27][9][5]

A rhododendron
Not described to species

Cf. Leucothoe

Undescribed[27]

A doghobble relative
Not described to species

Eucommiaceae

Eucommia

Eucommia montana[49][5]

Brown

A "hard rubber tree"

Eucommia montana
Fagaceae

Castaneophyllum

Undescribed[5]

A Castanea relative
Not described to species

Fagopsis

Fagopsis undulata[16]

(Knowlton) Wolfe & Wehr

A beech relative

Fagopsis undulata

Fagus

Fagus langevinii[50]

Manchester & Dillhoff

A beech

Fagus langevinii

Quercus

Undescribed[43][9][5]

An oak
Not described to species

Grossulariaceae Ribes

"Species 1"[27][9][5]

A current
Not described to species

"Species2"[27]

A current
Not described to species

Hamamelidaceae

Corylopsis

Corylopsis reedae[27][51]

Radtke, Pigg, & Wehr

A winter-hazel

Corylopsis reedae

Fothergilla

Fothergilla malloryi[51]

Radtke, Pigg, & Wehr

A witch alder

Fothergilla malloryi
Hydrangeaceae

Hydrangea

Undescribed[27]

A Hydrangea.
Not described to species

Philadelphus

Undescribed[27]

A mock-orange
Not described to species

Icacinaceae

Palaeophytocrene

Unidentified[43][9]

A Phytocrene relative
Not described to species

Iteaceae

Itea

Undescribed[16][5]

A virginia willow species
Not described to species.

Juglandaceae

Carya

Undescribed[9][5]

A walnut family relative.
Not described to species.

Cruciptera

Cruciptera simsonii[52]

(Brown) Manchester

A walnut family relative.

Juglans

Undescribed[9][5]

A walnut family relative.
Not described to species.

Pterocarya

Undescribed[43]

A wing nut
Not described to species

Lythraceae

Decodon

Undescribed[9]

A swamp loosestrife
Not described to species

Malvaceae

Craigia

Undescribed[43]

A Craigia species
Not described to species

Craigia sp.

Florissantia

Florissantia quilchenensis[53]

(Mathewes & Brooke) Manchester

A chocolate relative

Florissantia quilchenensis

Hibiscus

Undescribed[27]

A hibiscus
Not described to species

Plafkeria

Undescribed[43]

A linden relative
Not described to species

Tilia

Tilia johnsoni[16]

Wolfe & Wehr

A Linden

Tilia johnsoni

Cf. Tilia

Undescribed[48]

Linden relative fruits
Not described to species

Cf. Tilia fruits

Menispermaceae

Calycocarpum

Undescribed[24]

A moonseed
Not described to species

Moraceae

Morus

Undescribed[27]

A mulberry, two types known.
Not described to species

Myricaceae

Comptonia

Comptonia columbiana[16]

Dawson

A Comptonia

Comptonia columbiana

Nyssaceae

Tsukada

Tsukada davidiifolia[16]

Wolfe & Wehr

A dove-tree relative

Tsukada davidiifolia
Platanaceae

Langeranthus

Langeranthus dillhoffiorum[54]

Huegele & Manchester

A plane tree fruit taxon affiliated with Langeria

Langeria

Langeria magnifica[16]

Wolfe & Wehr

A plane tree
Formerly identified as a witch hazel relative

Langeria magnifica

Macginicarpa

Undescribed[43]

Manchester

A plane tree fruit taxon
Not described to species

Macginicarpa species

Macginitiea

Macginitiea gracilis[16]

(Lesquereux) Wolfe & Wehr

A plane tree relative

Macginitiea gracilis

Platananthus

Undescribed[48]

A sycamore stamen head
isolated stamen clusters placed as Macginistemon
Not described to species

Macginistemon stamen cluster

Polygalaceae

Deviacer

Undescribed[43]

A milkwort relative
Not described to species

Deviacer species

Ranunculaceae

Clematis

Undescribed[27]

A Clematis
Not described to species

Clematis species
Rosaceae

Amelanchier

Undescribed[27]

A service berry
Not described to species

Cf. Crataegus

Undescribed[27]

A hawthorn relative
Not described to species

Malus

Undescribed[5]

An apple
Not described to species

Cf. Malus

Undescribed[38]

A maloid species possibly apple or pear
Not described to species

Neviusia

Undescribed[27]

A snow-wreath
Not described to species

Oemleria

Oemleria janhartfordae[37]

Benedict, DeVore, & Pigg

An Osoberry

Photinia

Photinia pageae[16]

Wolfe & Wehr

A Christmas-berry relative

Photinia pagae

Aff. Physocarpus

Undescribed[9]

A possible nine-bark
Not described to species
Possibly stem Neillieae[39]

Prunus

Prunus cathybrownae[37]

Benedict, DeVore, & Pigg

A cherry relative

Prunus cathybrownae

"Species 1"[16]

A prunoid leaf
Not described to species

"Species 2"[16]

A prunoid leaf
Not described to species

"Species 3"[16]

A prunoid leaf
Not described to species

Pyracantha

Undescribed[38]

A firethorn sp.[38]
Tentative record, Not described to species.

Cf. Pyrus

Undescribed[38]

A maloid species possibly apple or pear
Not described to species

Rubus

Undescribed[27]

A blackberry
Not described to species

Aff. Sorbus

Undescribed[9][5]

A rowan relative
Not described to species.

Spiraea

Undescribed[9][5]

A bridal wreath
Not described to species.

Cf. Spiraea

Undescribed[38]

A bridal wreath relative
Not described to species

Sabiaceae

Meliosma

Undescribed[24]

A Meliosma species
Not described to species

Sabia

Undescribed[24]

A Sabia species
Not described to species

Salicaceae

Populus

Undescribed[9][5]

A cottonwood
Not described to species
First identified as †Populus lindgreni[17]

Pseudosalix

Undescribed[9]

A willow relative
Not described to species

Salix

Undescribed[9][5]

A willow
Not described to species

Sapindaceae

Acer?

"Acer" arcticum[55]

Heer, 1876

A possible maple

Acer

Acer hillsi[55]

Wolfe & Tanai

A maple

Acer republicense[55]

Wolfe & Tanai

A maple

Acer spitzi[55]

Wolfe & Tanai

A maple

Acer stonebergae[55]

Wolfe & Tanai

A maple

Acer toradense[55]

Wolfe & Tanai

A maple

Acer washingtonense[55]

Wolfe & Tanai

A maple

Acer wehri[55]

Wolfe & Tanai

A maple

Aesculus

Undescribed[9][56]

A horse chestnut
Not described to species.

Cf. Boniodendron

"Koelreuteria" arnoldii[16][57]

Becker

A sapindaceous species
first described as a Koelreuteria species,
considered Cf. Boniodendron by Wang et al. (2012).

"Koelreuteria" arnoldii

Bohlenia

Bohlenia americana[16]

(Brown) Wolf & Wehr

An extinct sapindalean species

Bohlenia americana

Dipteronia

Dipteronia brownii[58]

McClain & Manchester

A Dipteronia

Dipteronia brownii

Koelreuteria

Koelreuteria dilcheri[57]

Wang et al.

A Koelreuteria species

Koelreuteria dilcheri

Schisandraceae

Kadsura

Undescribed[2]

A kadsura species.
Not described to species

Schoepfiaceae

Schoepfia

Schoepfia republicensis[16]

(LaMotte) Wolfe & Wehr

A possible Schoepfia relative,
First described as "Cornus acuminata.

Schoepfia republicensis
Theaceae

Ternstroemites

"Species A"[16]

A theaceous species similar to Gordonia
Not described to species

Ternstroemites

"Species B"[16]

A theaceous species similar to Cleyera
Not described to species

Trochodendraceae?

Nordenskioldia

Undescribed[43]

A trochodendroid of uncertain placement.[42]
Fruits of the leaf taxon Zizyphoides
Not described to species.

Trochodendraceae

Paraconcavistylon

Paraconcavistylon wehrii[41][42]

(Manchester et al.)

A Trochodendrale
first described as "Concavistylon" wehrii
moved to a new genus in 2020.

Pentacentron

Pentacentron sternhartae[41]

Manchester et al.

A Trochodendrale

Pentacentron sternhartae

Tetracentron

Tetracentron hopkinsii[41]

Pigg et al.

A Trochodendrale,
possibly the leaves of Pentacentron sternhartae

Tetracentron hopkinsii

Trochodendron

Trochodendron nastae[59]

Pigg, Wehr, & Ickert-Bond

A Trochodendron
Possibly the leaves of Trochodendron drachukii

Trochodendron nastae

Zizyphoides

Undescribed[9]

A trochodendroid of uncertain placement.[42]
Leaves of the fruit taxon Nordenskioldia
Not described to species.

Zizyphoides species
Ulmaceae

Cedrelospermum

Undescribed[24]

An elm relative
Not described to species

Ulmus

Ulmus chuchuanus[60]

(Berry) LaMotte

An elm species
Leaves with features of Ulmus subg. Ulmus
fruits with features of Ulmus subg. Oreoptelea

Ulmus chuchuanus

Ulmus okanaganensis

Denk & Dillhoff

An elm species,
the fruits were first identified as Ulmus section Chaetoptelea.

Ulmus okanaganensis

Vitaceae

Vitis

Undescribed[27][43]

Grape seeds[43] and leaves[27]
Not described to species

Incertae sedis

Averrhoites

Undescribed[9]

A leaf morphotype of uncertain affiliation.
Not described to species.

Calycites

Calycites ardtunensis[43]

Crane

A winged fruit of unidentified affinities

Calycites ardtunensis

Pteroheterochrosperma

Pteroheterochrosperma horseflyensis[61]

Smith, Greenwalt & Manchester

A samara of uncertain affiliation.

Pteroheterochrosperma horseflyensis

Pteronepelys

Pteronepelys wehrii[62]

Manchester

A samara of uncertain affinities.

Pteronepelys wehrii

Republica

Republica hickeyi[16]

Wolfe & Wehr

An incertae sedis angiosperm
possibly of Hamamelididae affiliations

Republica hickeyi

Animals

Arthropods

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.[63] The most prevalent orders are Diptera and Hemiptera, each making up approximately 30% of the fossil insects known in 1992.

Blattodea

Family Genus species Authors Notes Images

Blaberidae

Undescribed

Undescribed[5]

A Diplopterine cockroach
Not described to genus/species

Blattoidae

Undescribed

Undescribed[63]

A blattoidean cockroach
Not described to genus/species

Undescribed Blattoidea

Isoptera

Undescribed

Undescribed[63]

Undescribed termites of uncertain affiliation

thumb|center|upright|undescribed isopteran

Coleoptera

A list of Coleopteran families identified by 1992 included Carabidae, Cerambycidae, Chrysomelidae, Curculionidae, Dytiscidae, Elateridae and Lucanidae,[63] but the beetle fauna has not been described in depth yet, with only two weevil species having been fully described.[64][65] A third beetle group belonging to the bean beetle tribe Pachymerini has been identified as palm beetles of the CaryobruchusSpeciomerus genus group.[66]

Family Genus Species Authors Notes Images
Brentidae

Eoceneithycerus

Eoceneithycerus carpenteri[64]

Legalov, 2013

An Ithycerinae weevil

Eoceneithycerus carpenteri

Ithyceroides

Ithyceroides klondikensis[65]

Legalov, 2015

An Ithycerinae weevil

Carabidae

Undescribed

Undescribed[63]

A ground beetle
Not described to genus/species

Unidentified Carabidae

Cerambycidae

Undescribed

Undescribed[63]

A long-horn beetle
Not described to genus/species

Chrysomelidae

CaryobruchusSpeciomerus genus group

Undescribed[66]

palm beetles in the tribe Pachymerini.
Not described to genus/species

Dytiscidae

Undescribed

Undescribed[63]

A diving beetle
Not described to genus/species

Elateridae

Undescribed

Undescribed[63]

A click beetle
Not described to genus/species

Lucanidae

Undescribed

Undescribed[63]

A stag beetle
Not described to genus/species

Unidentified

Undescribed

Undescribed[67]

A possible staphylinoid beetle
not described

Dermaptera

The order Dermaptera was first reported in 1992[63] 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.[68]

Family Genus Species Authors Notes Images
Forficulidae? Unidentified

"Forficulid species 1"[68]

A possible forficulid earwig with long cerci
Not described to genus/species

thumb|center|upright|"Forficulid species 1"

"Forficulid species 2"[68]

A possible forficulid earwig with short cerci
Not described to genus/species

"Forficulidae species 2"
undescribed

Diptera

Family Genus Species Authors Notes Images

Bibionidae

Undescribed

Undescribed[63]

A march fly
Not described to genus/species

thumb|center|upright|unidentified Bibionidae

Cecidomyiidae

Undescribed

Undescribed[69]

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

Cecidomyiidae gall on Prunus

Empididae

Undescribed

Undescribed[67]

A dagger fly
Not described to genus/species

Mycetophilidae

Undescribed

Undescribed[63]

A fungus gnat
Not described to genus/species

Pipunculidae

Metanephrocerus

Metanephrocerus belgardeae[70]

Archibald, Kehlmaier, & Mathewes, 2014

A pipunculid big-headed fly

Metanephrocerus belgardeae

Syrphidae

Unidentified

Unidentified[71]

A hover fly
Not described to genus/species

Tipulidae

Undescribed

Undescribed[63]

A crane fly
Not described to genus/species

Undescribed Tipulidae

Ephemeroptera

Lewis (1992) listed one species of Heptageniidae and three specimens that he did not place to family.[63] 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.[72] 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.[73]

Family Genus Species Authors Notes Images

Heptageniidae

Indeterminate

Indeterminate[63]

A flat headed mayfly nymph.
Tentatively suggested as Heptagenia or Stenonema by Lewis & Wehr (1993)
Deemed indeterminate below family level by Sinitchenkova (1999)[73]

Neoephemeridae

Neoephemera

Neoephemera antiqua[73]

Sinitchenkova, 1999

A squaregill mayfly

Hemiptera

Family Genus Species Authors Notes Images

Aphididae

Undescribed

Undescribed[63]

An aphid
Not described to genus/species.

Aphrophoridae

Aphrophora

Undescribed[63]

An aphrophorid spittlebug
Not described to species.

Aphrophora species

Petrolystra

Undescribed[63]

An aphrophorid spittlebug
Not described to species.

Cercopidae

Palecphora

Undescribed[63]

A cercopid froghopper
Not described to species.

Undescribed

Undescribed[63]

A cercopid froghopper
Not described to genus/species.

thumb|center|upright|undescribed Cercopidae

Fulgoroidea

Undescribed

Undescribed[63]

A frog hopper
Not described to genus/species.

Pentatomidae

Undescribed

Undescribed[63]

A Shield or stink bug
Not described to genus/species

thumb|center|upright|undescribed Pentatomidae

Hymenoptera

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.[74]

Family Genus Species Authors Notes Images
Angarosphecidae

Eosphecium

Undescribed[74]

An angarosphecid spheciform wasp.
Not described to species

Undescribed

Undescribed[74]

An angarosphecid spheciform wasp
Likely not Eosphecium.
Not described to species

Braconidae

Undescribed

Undescribed[63][74]

braconid parasitic wasps
Not described to genus/species.

thumb|center|upright|Unidentified Braconidae

Cimbicidae Leptostigma[75]

Leptostigma alaemacula[74][75]

Archibald & Rasnitsyn, 2023

A cenocimbicine cimbicid sawfly.

Leptostigma brevilatum[74][75]

Archibald & Rasnitsyn, 2023

A cenocimbicine cimbicid sawfly
Tentatively identified from Republic
described from the McAbee fossil beds

Leptostigma brevilatum?

Cynipidae

Undescribed

Undescribed[69]

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

Cynipidae gall on Prunus

Diapriidae

Undescribed

Undescribed[74]

A diapriid diaprioid wasp
Not described to species

Formicidae

Klondikia

Klondikia whiteae[76]

Dlussky & Rasnitsyn, 2003

An ant of uncertain subfamily affiliation

Myrmeciites

"Indesterminate"[77]

Archibald, Cover, & Moreau, 2006

A bulldog ant form genus

Myrmeciities sp.

Oecophylla

Oecophylla kraussei[78]

(Dlussky & Rasnitsyn, 1999)

An ant, described as Camponotites kraussei,
Moved to Oecophylla kraussei in 2017[79]

Propalosoma

Propalosoma gutierrezae[78]

Dlussky & Rasnitsyn, 1999

A bulldog ant, first described as a Rhopalosomatidae wasp,
moved to myrmeciinae in 2018[80]

Propalosoma gutierrezae

Undescribed

Undescribed[74]

Ants of uncertain subfamily placement.[74]

Unidentified formicidae

Halictidae (?)

Undescribed

Undescribed[74]

A possible sweat bee
Not described to genus/species

Ichneumonidae

Undescribed

Undescribed[74]

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

Undescribed Ichneumonidae

Megachilidae

Undescribed

Undescribed[74]

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

Pamphiliidae

Ulteramus

Ulteramus republicensis[81]

Archibald & Rasnitsyn, 2015

A parasitic wasp

Pompilidae

Undescribed

Undescribed[74]

A pompilid spider wasp
Not described to genus/species

Proctotrupidae

Undescribed

Undescribed[74]

A proctotrupid parasitic wasp
Not described to genus/species

Roproniidae

Undescribed

Undescribed[74]

A roproniid (sensu lato) proctotrupoid wasp
Not described to genus/species

Siricidae

Eourocerus

Eourocerus anguliterreus[82]

Archibald & Rasnitsyn, 2022

A siricine horntail.

Eourocerus anguliterreus

Scoliidae

Undescribed

Undescribed[74]

An archaeoscoliine scoliid wasps
Not described to genus/species

Sphecidae

Undescribed

Undescribed[74]

A sphecid (sensu stricto) wasp
Not described to genus/species

Tenthredinidae

Undescribed

Undescribed[74]

A Tenthredinid sawfly
Not described to genus/species

Vespidae

Undescribed

Undescribed[74]

A vespid wasp
Not described to genus/species

Lepidoptera

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.[83] Trace fossil evidence from leaf fossil herbivory indicates at least four other possible lepidopteran families were present in the formation. The

Family Genus Species Authors Notes Images

Coleophoridae

Undescribed

Undescribed[69]

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

Erebidae

Undescribed

Undescribed[83]

An arctiine tiger moth
Not described

Arctiinae
undescribed

Heliozelidae

Undescribed

Undescribed[69][84]

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

Incurvariidae

Aff. Incurvaria

Undescribed[69]

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

Nepticulidae

Stigmella

Undescribed[69]

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

Mecoptera

A number of mecopteran species belonging to the families Cimbrophlebiidae, Dinopanorpidae, Eorpidae, and Panorpidae are also known.[85]

Family Genus Species Authors Notes Images
Cimbrophlebiidae Cimbrophlebia

Cimbrophlebia brooksi[85]

Archibald, 2009

A Cimbrophlebiid scorpionfly

Cimbrophlebia brooksi

Cimbrophlebia westae[85]

Archibald, 2009

A Cimbrophlebiid scorpionfly

Cimbrophlebia westae
Dinopanorpidae Dinokanaga

Dinokanaga andersoni[86]

Archibald, 2005

A scorpion fly species

Dinokanaga andersoni

Dinokanaga dowsonae[86]

Archibald, 2005

A scorpion fly species

Dinokanaga dowsonae

Dinokanaga sternbergi[86]

Archibald, 2005

A scorpion fly species

Eorpidae Eorpa

Eorpa elverumi[87]

Archibald, Mathewes, & Greenwood, 2013

A mecopteran scorpionfly

Eorpa elverumi

?†Eorpa ypsipeda[87]

Archibald, Mathewes, & Greenwood, 2013

A mecopteran scorpionfly, tentatively identified

Possible E. ypsipeda[87]

Panorpidae

Undescribed

Undescribed[87]

Undescribed common scorpionflies
Not described to genus/species

Neuroptera

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.[88]

Family Genus Species Authors Year Notes Images

Chrysopidae

Adamsochrysa

Adamsochrysa wilsoni[89]

Makarkin & Archibald, 2013

2013

A nothochrysine green lacewing

Adamsochrysa wilsoni

Hemerobiidae

Archibaldia

Archibaldia wehri[90][91][92]

(Makarkin, Archibald, & Oswald, 2003)

2003

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

Ithonidae

Allorapisma

Allorapisma chuorum[93]

Makarkin & Archibald, 2009

2009

A moth lacewing

Allorapisma chuorum
Palaeopsychops

Palaeopsychops marringerae[94]

Archibald & Makarkin, 2006

2006

A polystechotid group[95] moth lacewing

Palaeopsychops marringerae

Palaeopsychops timmi[94]

Archibald & Makarkin, 2006

2006

A polystechotid group[95] moth lacewing

Palaeopsychops timmi
Polystoechotites

Polystoechotites barksdalae[94]

Archibald & Makarkin, 2006

2006

A polystechotid group[95] moth lacewing

Polystoechotites barksdalae

Polystoechotites falcatus[94]

Archibald & Makarkin, 2006

2006

A polystechotid group[95] moth lacewing

Polystoechotites falcatus

Polystoechotites lewisi[94]

Archibald & Makarkin, 2006

2006

A polystechotid group[95] moth lacewing

Polystoechotites lewisi

Nymphidae

?Nymphes

Nymphes? georgei[96]

Archibald, Makarkin, & Ansorge, 2009

2009

A nymphid lacewing, possibly a species of †Epinesydrion[97]

Osmylidae

Osmylidia

Osmylidia glastrai[98]

Makarkin, Archibald, & Mathewes, 2021

2021

A protosmyline osmylid lacewing

Psychopsidae?

Ainigmapsychops

Ainigmapsychops inexspectatus[88]

Makarkin & Archibald, 2014

2014

A possible psychopsid lacewing

Ainigmapsychops inexspectatus

Odonata

Family Genus Species Authors Year Notes Images
Aeshnidae

Antiquiala

Antiquiala snyderae[99]

Archibald & Cannings, 2019

2019

A darner dragonfly

Antiquiala snyderae

Idemlinea

Idemlinea versatilis[99]

Archibald & Cannings, 2019

2019

A darner dragonfly

Idemlinea versatilis

Ypshna

Ypshna brownleei[99]

Archibald & Cannings, 2019

2019

A darner dragonfly

Ypshna brownleei
†Dysagrionidae

Dysagrion

Dysagrion pruettae[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Dysagrion pruettae

Dysagrionites

Dysagrionites delinei[100]

Archibald & Cannings, 2021

2021

A dysagrionine cephalozygopteran odonate

Dysagrionites delinei
Okanagrion

Okanagrion dorrellae[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Okanagrion dorrellae

Okanagrion hobani[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Okanagrion hobani

Okanagrion liquetoalatum[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Okanagrion liquetoalatum

Okanagrion threadgillae[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Okanagrion threadgillae

Okanagrion worleyae[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Okanagrion worleyae

Okanopteryx

Okanopteryx jeppesenorum[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Okanopteryx jeppesenorum

Stenodiafanus

Stenodiafanus westersidei[100]

Archibald & Cannings, 2021

2021

A Dysagrionine cephalozygopteran odonate

Stenodiafanus westersidei

Euphaeidae

Republica

Republica weatbrooki[101]

Archibald & Cannings, 2021

2021

A gossamerwing damselfly.
Not to be confused with the plant Republica,
also from the formation

Republica weatbrooki

†Whetwhetaksidae

Whetwhetaksa

Whetwhetaksa millerae[100]

Archibald & Cannings

2021

A cephalozygopteran odonate

Whetwhetaksa millerae

Orthoptera

Family Genus Species Authors Year Notes Images

Palaeorehniidae

Republicopteron

Republicopteron douseae[102]

Archibald, Gu, & Mathewes

2022

A grasshopper/hump-back grig relative

Republicopteron douseae

Phasmatodea

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.[103] 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.[104]

Family Genus Species Authors Year Notes Images

Susumaniidae

Eoprephasma

Eoprephasma hichensi[103]

Archibald & Bradler, 2015

2015

A Susumaniinae stick insect species

Raphidioptera

Family Genus Species Authors Year Notes Images

Raphidiidae

Megaraphidia

Megaraphidia klondika[105]

Archibald & Makarkin, 2021

2021

A raphidiid snakefly

Trichoptera

Trichopterans are known mainly from laraval cases and occasional isolated wings.[106]

Family Genus species Authors Year Notes Images

Phryganeidae

Unidentified

Unidentified[63][106]

giant caddisflies
Not described to genus/species

Limnephilidae

Unidentified

unidentified[63]

northern caddisflies
Not described to genus/species

Vertebrates

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.[107] 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.[108] Research tapered off until a series of fish were collected in the Toroda Creek Graben northwest of Republic by R. C. Pearson during his compilation of the Geologic map of the Bodie Mountain quadrangle, Ferry and Okanogan Counties, Washington. The fossils were tentatively identified by paleoichthyologist David Dunkle as members of the genera Amyzon, Tricophanes, Erismatopterus and an undefined salmonid.[109] Pearson sent almost all of the specimens collected to the Smithsonian, but the fossils were never accessioned into the collections there 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.[110] 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.[111][112] 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.[109] A year later the percopsid fossils were also described as Libotonius pearsoni, extending the range of the genus south from the Allenby Formation.[111]

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.

Family Genus Species Authors Year Notes Images

Amiidae

Amia

"Amia" hesperia

Wilson, 1977

1977

A bowfin, known from isolated scales

"Amia" hesperia scale

Catostomidae

Amyzon

Amyzon aggregatum

Wilson, 1977

1977

A sucker

Amyzon aggregatum

Catostomidae

Amyzon

Unidentified[113]

(Wilson, 1977)

1977

A sucker, originally identified as Amyzon aggregatum

Salmonidae

Eosalmo

Eosalmo driftwoodensis[114]

Wilson, 1977

1999

A Salmon

Eosalmo driftwoodensis

Hiodontidae

Hiodon

Hiodon woodruffi[109]

Wilson, 1978

1978

A mooneye, first described as "Eohiodon" woodruffi.[109]

Hiodon woodruffi

†Libotoniidae

Libotonius

Libotonius pearsoni[111]

Wilson, 1979

1979

A sand roller relative.

Libotonius pearsoni

incertae sedis (Aves)

Unidentified

"Unnamed"[115]

2019

indeterminate feathers and a skeleton

Unidentified feather

References

  1. 1.0 1.1 Dillhoff, R.M.; Dillhoff, T.A.; Greenwood, D.R.; DeVore, M.L.; Pigg, K.B. (2013). "The Eocene Thomas Ranch flora, Allenby Formation, Princeton, British Columbia, Canada". Botany 91 (8): 514–529. doi:10.1139/cjb-2012-0313. 
  2. 2.0 2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 2.9 Wehr, W.. "Middle Eocene insects and plants of the Okanogan Highlands". Contributions to the Paleontology and Geology of the West Coast. Seattle, Washington: University of Washington Press. pp. 99–109. 
  3. 3.0 3.1 Pigg, K. B.; DeVore, M. L.; Greenwood, D. R.; Sundue, M. A.; Schwartsburd, P.; Basinger, J. F. (2021). "Fossil Dennstaedtiaceae and Hymenophyllaceae from the Early Eocene of the Pacific Northwest". International Journal of Plant Sciences 182 (9): 793–807. doi:10.1086/715633. 
  4. Joseph, N. L. (1988). "Important Eocene Flora and Fauna Unearthed at Republic, Washington". Rocks & Minerals 63 (2): 146–151. doi:10.1080/00357529.1988.11761830. Bibcode1988RoMin..63..146J. 
  5. 5.00 5.01 5.02 5.03 5.04 5.05 5.06 5.07 5.08 5.09 5.10 5.11 5.12 5.13 5.14 5.15 5.16 5.17 5.18 Greenwood, D.R.; Archibald, S.B.; Mathewes, R.W; Moss, P.T. (2005). "Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape". Canadian Journal of Earth Sciences 42 (2): 167–185. doi:10.1139/e04-100. Bibcode2005CaJES..42..167G. 
  6. Arnold, C. A. (1955). "A Tertiary Azolla from British Columbia". Contributions from the Museum of Paleontology, University of Michigan 12 (4): 37–45. http://deepblue.lib.umich.edu/bitstream/2027.42/48289/2/ID129.pdf. 
  7. 7.0 7.1 Greenwood, D.R.; Pigg, K.B.; Basinger, J.F.; DeVore, M.L. (2016). "A review of paleobotanical studies of the Early Eocene Okanagan (Okanogan) Highlands floras of British Columbia, Canada, and Washington, U.S.A.". Canadian Journal of Earth Sciences 53 (6): 548–564. doi:10.1139/cjes-2015-0177. Bibcode2016CaJES..53..548G. 
  8. Hopkins, D.; Johnson, K. (1997). "First Record of cycad leaves from the Eocene Republic flora". Washington Geology 25 (4): 37. https://www.dnr.wa.gov/Publications/ger_washington_geology_1997_v25_no4.pdf. Retrieved 29 September 2021. 
  9. 9.00 9.01 9.02 9.03 9.04 9.05 9.06 9.07 9.08 9.09 9.10 9.11 9.12 9.13 9.14 9.15 9.16 9.17 9.18 9.19 9.20 Dillhoff, R.M.; Leopold, E.B.; Manchester, S.R. (2005). "The McAbee flora of British Columbia and its relations to the Early-Middle Eocene Okanagan Highlands flora of the Pacific Northwest". Canadian Journal of Earth Sciences 42 (2): 151–166. doi:10.1139/e04-084. Bibcode2005CaJES..42..151D. http://www.fossilmuseum.net/Fossil_Sites/mcabee/mcabeefossils.pdf. 
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