Paleobiota of the Klondike Mountain Formation

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	Dennstaedtia christophelii
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	Metasequoia occidentalis
	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 .	Nuphar carlquistii seeds

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
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]	"Schoepfia" republicensis
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
Oxalidaceae?	† Averrhoites	Undescribed [9]		A leaf morphotype of uncertain affiliation. First described as visually similar to Averrhoa Not described to species.
Platanaceae	† Langeranthus	† Langeranthus dillhoffiorum [56]	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 [57]	Heer, 1876	A possible maple
	Acer	† Acer hillsi [57]	Wolfe & Tanai	A maple
		† Acer republicense [57]	Wolfe & Tanai	A maple
		† Acer spitzi [57]	Wolfe & Tanai	A maple
		† Acer stonebergae [57]	Wolfe & Tanai	A maple
		† Acer toradense [57]	Wolfe & Tanai	A maple
		† Acer washingtonense [57]	Wolfe & Tanai	A maple
		† Acer wehri [57]	Wolfe & Tanai	A maple
	Aesculus	Undescribed [9] [58]		A horse chestnut Not described to species.
	Cf. Boniodendron	† "Koelreuteria" arnoldii [16] [59]	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 [60]	McClain & Manchester	A Dipteronia	Dipteronia brownii
	Koelreuteria	† Koelreuteria dilcheri [59]	Wang et al.	A Koelreuteria species	Koelreuteria dilcheri
Schisandraceae	Kadsura	Undescribed [2]		A kadsura species. Not described to species
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 [61]	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 [62]	(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	† Calycites	† Calycites ardtunensis [43]	Crane	A winged fruit of unidentified affinities	Calycites ardtunensis
	† Pteroheterochrosperma	† Pteroheterochrosperma horseflyensis [63]	Smith, Greenwalt & Manchester	A samara of uncertain affiliation.	Pteroheterochrosperma horseflyensis
	† Pteronepelys	† Pteronepelys wehrii [64]	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. ^[65] 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 [65]		A blattoidean cockroach Not described to genus/species	Undescribed Blattoidea
Isoptera	Undescribed	Undescribed [65]		Undescribed termites of uncertain affiliation	undescribed isopteran

Coleoptera

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]

Family	Genus	Species	Authors	Notes	Images
Brentidae	† Eoceneithycerus	† Eoceneithycerus carpenteri [66]	Legalov, 2013	An Ithycerinae weevil	Eoceneithycerus carpenteri
	† Ithyceroides	† Ithyceroides klondikensis [67]	Legalov, 2015	An Ithycerinae weevil
Carabidae	Undescribed	Undescribed [65]		A ground beetle Not described to genus/species	Unidentified Carabidae
Cerambycidae	Undescribed	Undescribed [65]		A long-horn beetle Not described to genus/species
Chrysomelidae	Caryobruchus – Speciomerus genus group	Undescribed [68]		palm beetles in the tribe Pachymerini. Not described to genus/species
Dytiscidae	Undescribed	Undescribed [65]		A diving beetle Not described to genus/species
Elateridae	Undescribed	Undescribed [65]		A click beetle Not described to genus/species
Lucanidae	Undescribed	Undescribed [65]		A stag beetle Not described to genus/species
Unidentified	Undescribed	Undescribed [69]		A possible staphylinoid beetle not described

Dermaptera

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]

Family	Genus	Species	Authors	Notes	Images
Forficulidae?	Unidentified	"Forficulid species 1" [70]		A possible forficulid earwig with long cerci Not described to genus/species	"Forficulid species 1"
		"Forficulid species 2" [70]		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 [65]		A march fly Not described to genus/species	unidentified Bibionidae
Cecidomyiidae	Undescribed	Undescribed [71]		Trace fossils Cecidomyiid midge galling on various host leaves Not described to genus/species	Cecidomyiidae gall on Prunus
Empididae	Undescribed	Undescribed [69]		A dagger fly Not described to genus/species
Mycetophilidae	Undescribed	Undescribed [65]		A fungus gnat Not described to genus/species
Pipunculidae	† Metanephrocerus	† Metanephrocerus belgardeae [72]	Archibald, Kehlmaier, & Mathewes, 2014	A pipunculid big-headed fly	Metanephrocerus belgardeae
Syrphidae	Unidentified	Unidentified [73]		A hover fly Not described to genus/species
Tipulidae	Undescribed	Undescribed [65]		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. ^[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]

Family	Genus	Species	Authors	Notes	Images
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]	Heptageniidae nymph
Neoephemeridae	Neoephemera	† Neoephemera antiqua [75]	Sinitchenkova, 1999	A squaregill mayfly

Hemiptera

Family	Genus	Species	Authors	Notes	Images
Aphididae	Undescribed	Undescribed [65]		An aphid Not described to genus/species.
Aphrophoridae	Aphrophora	Undescribed [65]		An aphrophorid spittlebug Not described to species.	Aphrophora species
	† Petrolystra	Undescribed [65]		An aphrophorid spittlebug Not described to species.
Cercopidae	† Palecphora	Undescribed [65]		A cercopid froghopper Not described to species.
	Undescribed	Undescribed [65]		A cercopid froghopper Not described to genus/species.	undescribed Cercopidae
Fulgoroidea	Undescribed	Undescribed [65]		A frog hopper Not described to genus/species.
Pentatomidae	Undescribed	Undescribed [65]		A Shield or stink bug Not described to genus/species	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. ^[76]

Family	Genus	Species	Authors	Notes	Images
Angarosphecidae	† Eosphecium	Undescribed [76]		An angarosphecid spheciform wasp. Not described to species
	Undescribed	Undescribed [76]		An angarosphecid spheciform wasp Likely not Eosphecium . Not described to species
Braconidae	Undescribed	Undescribed [65] [76]		braconid parasitic wasps Not described to genus/species.	Unidentified Braconidae
Cimbicidae	† Leptostigma [77]	† Leptostigma alaemacula [76] [77]	Archibald & Rasnitsyn, 2023	A cenocimbicine cimbicid sawfly.
		† Leptostigma brevilatum [76] [77]	Archibald & Rasnitsyn, 2023	A cenocimbicine cimbicid sawfly Tentatively identified from Republic described from the McAbee fossil beds	Leptostigma brevilatum ?
Cynipidae	Undescribed	Undescribed [71]		Trace fossils cynipid Cynipoid gallwasp galling on various host leaves Not described to genus/species	Cynipidae gall on Prunus
Diapriidae	Undescribed	Undescribed [76]		A diapriid diaprioid wasp Not described to species
Formicidae	† 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	Myrmeciities sp.
	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]	Propalosoma gutierrezae
	Undescribed	Undescribed [76]		Ants of uncertain subfamily placement. [76]	Unidentified formicidae
Halictidae (?)	Undescribed	Undescribed [76]		A possible sweat bee Not described to genus/species
Ichneumonidae	Undescribed	Undescribed [76]		ichneumonid parasitic wasps unplaced to subfamily Not described to genus/species	Undescribed Ichneumonidae
Megachilidae	Undescribed	Undescribed [76]		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	Undescribed	Undescribed [76]		A pompilid spider wasp Not described to genus/species
Proctotrupidae	Undescribed	Undescribed [76]		A proctotrupid parasitic wasp Not described to genus/species
Roproniidae	Undescribed	Undescribed [76]		A roproniid (sensu lato) proctotrupoid wasp Not described to genus/species
Siricidae	† Eourocerus	† Eourocerus anguliterreus [84]	Archibald & Rasnitsyn, 2022	A siricine horntail.	Eourocerus anguliterreus
Scoliidae	Undescribed	Undescribed [76]		An archaeoscoliine scoliid wasps Not described to genus/species
Sphecidae	Undescribed	Undescribed [76]		A sphecid (sensu stricto) wasp Not described to genus/species
Tenthredinidae	Undescribed	Undescribed [76]		A Tenthredinid sawfly Not described to genus/species
Vespidae	Undescribed	Undescribed [76]		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. ^[85] 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 [71]		Trace fossils Coleophorid hole feeding and larval cases Not described to genus/species
Erebidae	Undescribed	Undescribed [85]		An arctiine tiger moth Not described	Arctiinae undescribed
Heliozelidae	Undescribed	Undescribed [71] [86]		Trace fossils heliozelid leaf mining similar to Antispila mines Not described to genus/species
Incurvariidae	Aff. Incurvaria	Undescribed [71]		Trace fossils incurvariid leaf mining similar to Incurvaria Not described to genus/species
Nepticulidae	Stigmella	Undescribed [71]		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. ^[87]

Family	Genus	Species	Authors	Notes	Images
†Cimbrophlebiidae	† Cimbrophlebia	† Cimbrophlebia brooksi [87]	Archibald, 2009	A Cimbrophlebiid scorpionfly	Cimbrophlebia brooksi
		† Cimbrophlebia westae [87]	Archibald, 2009	A Cimbrophlebiid scorpionfly	Cimbrophlebia westae
†Dinopanorpidae	† Dinokanaga	† Dinokanaga andersoni [88]	Archibald, 2005	A scorpion fly species	Dinokanaga andersoni
		† Dinokanaga dowsonae [88]	Archibald, 2005	A scorpion fly species	Dinokanaga dowsonae
		† Dinokanaga sternbergi [88]	Archibald, 2005	A scorpion fly species
†Eorpidae	† Eorpa	† Eorpa elverumi [89]	Archibald, Mathewes, & Greenwood, 2013	A mecopteran scorpionfly	Eorpa elverumi
		?† Eorpa ypsipeda [89]	Archibald, Mathewes, & Greenwood, 2013	A mecopteran scorpionfly, tentatively identified	Possible E. ypsipeda
Panorpidae	Undescribed	Undescribed [89]		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. ^[90]

Family	Genus	Species	Authors	Year	Notes	Images
Chrysopidae	† Adamsochrysa	† Adamsochrysa wilsoni [91]	Makarkin & Archibald, 2013	2013	A nothochrysine green lacewing	Adamsochrysa wilsoni
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)
Ithonidae	† Allorapisma	† Allorapisma chuorum [95]	Makarkin & Archibald, 2009	2009	A moth lacewing	Allorapisma chuorum
	† Palaeopsychops	† Palaeopsychops marringerae [96]	Archibald & Makarkin, 2006	2006	A polystechotid group [97] moth lacewing	Palaeopsychops marringerae
		† Palaeopsychops timmi [96]	Archibald & Makarkin, 2006	2006	A polystechotid group [97] moth lacewing	Palaeopsychops timmi
	† Polystoechotites	† Polystoechotites barksdalae [96]	Archibald & Makarkin, 2006	2006	A polystechotid group [97] moth lacewing	Polystoechotites barksdalae
		† Polystoechotites falcatus [96]	Archibald & Makarkin, 2006	2006	A polystechotid group [97] moth lacewing	Polystoechotites falcatus
		† Polystoechotites lewisi [96]	Archibald & Makarkin, 2006	2006	A polystechotid group [97] moth lacewing	Polystoechotites lewisi
Nymphidae	? Nymphes	† Nymphes? georgei [98]	Archibald, Makarkin, & Ansorge, 2009	2009	A nymphid lacewing, possibly a species of † Epinesydrion [99]	Nymphes georgei
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	Ainigmapsychops inexspectatus

Odonata

Family	Genus	Species	Authors	Year	Notes	Images
Aeshnidae	† Antiquiala	† Antiquiala snyderae [101]	Archibald & Cannings, 2019	2019	A darner dragonfly	Antiquiala snyderae
	† Idemlinea	† Idemlinea versatilis [101]	Archibald & Cannings, 2019	2019	A darner dragonfly	Idemlinea versatilis
	† Ypshna	† Ypshna brownleei [101]	Archibald & Cannings, 2019	2019	A darner dragonfly	Ypshna brownleei
†Dysagrionidae	† Dysagrion	† Dysagrion pruettae [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Dysagrion pruettae
	† Dysagrionites	† Dysagrionites delinei [102]	Archibald & Cannings, 2021	2021	A dysagrionine cephalozygopteran odonate	Dysagrionites delinei
	† Okanagrion	† Okanagrion dorrellae [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Okanagrion dorrellae
		† Okanagrion hobani [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Okanagrion hobani
		† Okanagrion liquetoalatum [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Okanagrion liquetoalatum
		† Okanagrion threadgillae [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Okanagrion threadgillae
		† Okanagrion worleyae [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Okanagrion worleyae
	† Okanopteryx	† Okanopteryx jeppesenorum [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Okanopteryx jeppesenorum
	† Stenodiafanus	† Stenodiafanus westersidei [102]	Archibald & Cannings, 2021	2021	A Dysagrionine cephalozygopteran odonate	Stenodiafanus westersidei
Euphaeidae	† Republica	† Republica weatbrooki [103]	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 [102]	Archibald & Cannings	2021	A cephalozygopteran odonate	Whetwhetaksa millerae

Orthoptera

Family	Genus	Species	Authors	Year	Notes	Images
Palaeorehniidae	† Republicopteron	† Republicopteron douseae [104]	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. ^[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]

Family	Genus	Species	Authors	Year	Notes	Images
†Susumaniidae	† Eoprephasma	† Eoprephasma hichensi [105]	Archibald & Bradler, 2015	2015	A Susumaniinae stick insect species

Raphidioptera

Family	Genus	Species	Authors	Year	Notes	Images
Raphidiidae	† Megaraphidia	† Megaraphidia klondika [107]	Archibald & Makarkin, 2021	2021	A raphidiid snakefly

Trichoptera

Trichopterans are known mainly from laraval cases and occasional isolated wings. ^[108]

Family	Genus	species	Authors	Year	Notes	Images
Phryganeidae	Unidentified	Unidentified [65] [108]			giant caddisflies Not described to genus/species
Limnephilidae	Unidentified	unidentified [65]			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. ^[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 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. ^[111] 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. ^[112] 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. ^{[113] [114]} 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. ^[111] A year later the percopsid fossils were also described as Libotonius pearsoni, extending the range of the genus south from the Allenby Formation. ^[113]

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 [115]	(Wilson, 1977)	1977	A sucker, originally identified as Amyzon aggregatum
Salmonidae	† Eosalmo	† Eosalmo driftwoodensis [116]	Wilson, 1977	1999	A Salmon	Eosalmo driftwoodensis
Hiodontidae	Hiodon	† Hiodon woodruffi [111]	Wilson, 1978	1978	A mooneye, first described as "Eohiodon" woodruffi. [111]	Hiodon woodruffi
†Libotoniidae	† Libotonius	† Libotonius pearsoni [113]	Wilson, 1979	1979	A sand roller relative.	Libotonius pearsoni
incertae sedis (Aves)	Unidentified	"Unnamed" [117]		2019	indeterminate feathers and a skeleton	Unidentified feather

References

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