Klondike Mountain Formation

Klondike Mountain Formation
Stratigraphic range: Ypresian
50.1–48.9 Ma

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Klondike Mountain Formation; Stratigraphic range: Ypresian; 50.1–48.9 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Klondike Mountain Formation outcrop at site A0307
Type	Geological formation
Unit of
Sub-units	Tom Thumb Tuff, Lower & Upper Members
Underlies	Glacial deposits
Overlies	& others
Thickness	up to 3,200 ft (980 m)
Lithology
Primary	Variable
Other	See text
Location
Coordinates	48°42′N 118°42′W / 48.7°N 118.7°WCoordinates: 48°42′N 118°42′W / 48.7°N 118.7°W
Approximate paleocoordinates	54°24′N 101°54′W / 54.4°N 101.9°W
Region	Okanagan highlands
Country	United States
Extent	Okanagan highlands
Type section
Named for
Named by	Muessig
Year defined	1962
	Klondike Mountain Formation (the United States)Show map of the United StatesShow map of Washington (state)Show all

The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington state. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Washington, is composed of volcanic rocks in the upper unit and volcanics plus lacustrine (lakebed) sedimentation in which a lagerstätte with exceptionally well-preserved plant and insect fossils has been found, along with fossil epithermal hot springs.

The formation is the youngest in a group of formations which belong to the Challis Sequence rocks. The formation unconformably overlies rocks of the Eocene and much older Triassic and Permian formations. The formation is bounded on its edges by a series of high-angle strike slip faults, which have contained the Klondike Mountain Formation in a series of graben structures, such as the Republic Graben. Public access to a fossiliferous outcrop at the north end of Republic is mediated by the nonprofit Stonerose Interpretive Center.

Extent[]

The formation is located in northern Ferry County, Washington, with the majority of the sedimentation in the Republic and Curlew Basins on the east and in the Toroda Creek area to the north west. The town of Republic, Washington is situated at the southern end of the formation, with outcrops within the city itself. The formation was named for Klondike Mountain which is just to the northeast of the city, and where the type locality is located.^[1] The Curlew basin is situated north of Republic, with the northern edge along the Kettle River and the community of Curlew, Washington near the northeastern edge.^[2]^[3]

The formation is the southernmost of a string of preserved Eocene highland lakebeds in Washington state and British Columbia.^[4] The lake system, within the Okanagan highlands, extends from the Klondike Mountain Formation north approximately 1,000 kilometres (1,000,000 m) in to southern central British Columbia.^[5]

Age[]

Early dating of the formation was based primarily on identification and correlation of the fossils found in the Tom Thumb Tuff, with Joseph Umpleby (1910) reporting a putative age of Early Miocene.^[6] This date was based in examination of fossils by C. R. Eastman, who thought them to be similar to those found in the Florissant Formation of Colorado, which at the time was also considered Miocene. This age was retained by Edward W. Berry (1929), who included the Klondike Mountain Formation fossil lakebeds as part of the Latah Formation.^[7] The age of the Formation has been revised in the following hundred years, with Roland W. Brown (1936) identifying the deposits as being older than the Latah Formation.^[8] In a later written communication circa 1958, Brown again revised the age still older, stating the fossils found in the area of Mount Elizabeth indicated an Oligocene age. This age was used by Parker and Calkins (1964) for work on the Curlew Quadrangle of Ferry County.^[9] Work by Pearson and Obradovich (1977) refined the Sanpoil Volcanics age to between 52 to 51 million years ago, and a general age estimate for the Klondike Mountain Formation between 50.3 ± 1.7 million years ago and 42.3 ± 2.0 million years ago.^[10] Since then the fossil-bearing strata of the Formation have been radiometrically dated, to give an estimate of the Ypresian, the mid stage of the early Eocene,49.4 ± .5 million years ago,^[5] which was revised to an oldest age estimate of 51.2 ± 0.1 million years ago which given based on detrital zircon isotopic data published in 2021.^[11]

Lithology[]

Parker and Calkins (1964) noted the association of the Klondike Mountain Formation with the gold and silver deposits of the Republic District and suggested it as a potential host to more ore deposits in the Curlew Quadrangle.^[9] The epithermal gold deposits occurring in the Sanpoil volcanics terminate directly below the unconformity where the volcanics contact the base of the Klondike Mountain Formation^[12] or sometimes penetrate into the Formation's lowest unit.^[13] Hydrothermal sinter deposits are known from the lowest portions of the Formation and are thought to represent hydrothermal eruption areas.^[13] In general the lower portions of the Formation have a large amount of hydrothermal alteration, and areas around vents are rich in pyrite and silica. two products of natural hydrothermic sintering. The areas above that show a transition to mudstones, siltstones and sandstones grading from fine-grained material into coarser materials moving up the strata column. The finely-bedded stones show the greatest numbers of fossils and the finest preservation of details.^[14]

Paleoenvironment and paleoclimate[]

The lake bed sediments preserve a diverse array of plants, insects, and fishes, notably the biota called the Republic flora.^[15] The Okanagan lake system, which includes the Klondike Mountain Formation, has been classified as one of the great Canadian lagerstätten.^[16] The area likely had a mesic upper microthermal to lower mesothermal climate, in which winter temperatures rarely dropped low enough for snow, and which were seasonably equitable.^[17]

The Okanagan highlands paleoforest surrounding the lakes have been described as precursors the to modern Temperate broadleaf and mixed forests of Eastern North America and Eastern Asia. Based on the fossil biotas the lakes were higher and cooler then the coeval coastal forests preserved in the Puget Group and Chuckanut Formation of Western Washington, which are described as lowland tropical forest ecosystems. Estimates of the paleoelevation between 0.7–1.2 km (0.43–0.75 mi) higher than the coastal forests. This is consistent with the paleoelevation estimates for the lake systems, which range between 1.1–2.9 km (1,100–2,900 m), which is similar to the modern elevation 0.8 km (0.50 mi), but higher.^[17]

Estimates of the mean annual temperature have been derived from Climate leaf analysis multivariate program (CLAMP) and leaf margin analysis (LMA) of the Republic paleoflora. The CLAMP results after multiple linear regressions for Republic gave a mean annual temperature of approximately 8.0 °C (46.4 °F), while the LMA gave 9.2 ± 2.0 °C (48.6 ± 3.6 °F). This is lower than the mean annual temperature estimates given for the coastal Puget Group, which is estimated to have been between 15–18.6 °C (59.0–65.5 °F). The bioclimactic analysis for Republic suggests a mean annual precipitation amount of 115 ± 39 cm (45 ± 15 in).^[17]

Paleobiota[]

The formation is host to a highly diverse assemblage of plants and insects including some of the last records of taxa before extinction and the first appearances of taxa.

Lycophytes[]

Family	Genus	Species	Authors	Year	Notes	Images
Selaginellaceae	Selaginella	Undescribed^[18]		A spikemoss Not described to species

Pteridophytes[]

Family	Genus	Species	Authors	Notes	Images
Cystopteridaceae	Cf. Cystopteris	Undescribed^[18]		A possible bladder fern relative Not described to species
Equisetaceae	Equisetum	Undescribed^[19]^[18]		A scouring rush Not described to species
Lygodiaceae	Lygodium	Undescribed^[18]^[17]		A climbing fern Not described to species
Salviniaceae	Azolla	†Azolla primaeva^[20]	Arnold	A mosquito fern	Azolla primaeva
Salviniaceae	Salvinia	Undescribed^[21]		A "watermoss" species. Not described to species Found only in Curlew half graben sites.

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^[22]^[23]		A zamiaceous cycad. Not described to genus/species	Undescribed zamiaceous leaf

Gingkophytes[]

Family	Genus	Species	Authors	Notes	Images
Ginkgoaceae	Ginkgo	†Ginkgo biloba^[24]	Linnaeus	A ginkgo	Ginkgo biloba
Ginkgoaceae	Ginkgo	†Ginkgo dissecta^[24]		A ginkgo	Ginkgo dissecta

Pinophytes[]

Family	Genus	Species	Authors	Notes	Images
Cupressaceae	Calocedrus	Undescribed^[23]^[25]		An incense cedar Not described to species
Cupressaceae	Chamaecyparis	Undescribed^[26]		A false cypress Not described to species Possibly in the Callitropsis nootkatensis lineage.^[27]
Cupressaceae	Cryptomeria	Undescribed^[26]^[17]		A sugi Not described to species
Cupressaceae	Glyptostrobus	Undescribed^[26]		A Chinese swamp cypress Not described to species
Cupressaceae	Metasequoia	†Metasequoia occidentalis^[28]^[15]	(Newberry) Chaney	A dawn redwood	Metasequoia occidentalis
Cupressaceae	Sequoia	†Sequoia affinis^[28]	Lesquereux	A coast redwood Reported as "Sequioa langsdorfii" by Brown, 1935	Sequoia affinis
Cupressaceae	Taiwania	Undescribed^[21]		A Taiwania species not described to species
Cupressaceae	Taxodium	†Taxodium dubium^[7]^[28]	(Sternberg) Heer	A bald cypress
Cupressaceae	Thuja	Undescribed^[26]		An arborvitae Not described to species
Pinaceae	Abies	†Abies milleri^[29]	& Wehr	Oldest true fir described	Abies milleri
Pinaceae	Picea	Undescribed^[26]		A spruce Not described to species
Pinaceae	Pinus	†Pinus latahensis^[7]	Berry	A 5 needle pine	Pinus latahensis
Pinaceae	Pinus	†^[7]	Berry	A 3 needle pine, jr homonym to Pinus macrophylla Lindley 1839
Pinaceae	Pinus	†^[7]	Berry	A pine seed morphogenus
Pinaceae	Pinus	†^[7]	Berry	A possible 4 needled pine Noted by Berry as "highly improbable that this should represent a distinct botanic species"
Pinaceae	Pseudolarix	†Pseudolarix wehrii^[30]	Gooch	A golden larch Originally described as Pseudolarix americana.^[30]	Pseudolarix wehrii
Pinaceae	Tsuga	Undescribed^[26]		A hemlock Not described to species
	Sciadopitys	Undescribed^[26]		An umbrella pine Not described to species	Sciadopitys species
Taxaceae	Amentotaxus	Undescribed^[26]		A yew Not described to species	Amentotaxus sp.
Taxaceae	Cephalotaxus	Undescribed^[26]		A yew Originally described placed in the Miocene ^[7]^[31] Not described to species
Taxaceae	Taxus	Undescribed^[26]		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.^[32] 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.,^[33] though a second member, , has been described from the Princeton Chert.^[34] Wehr (1995) illustrated two fossils that were tentatively identified as fruits of the banana genus Ensete and the extinct myrtle genus respectively,^[35] 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.^[33]

Family	Genus	Species	Authors	Notes	Images
Nymphaeaceae	Nuphar	†Nuphar carlquistii^[33]	DeVore, Taylor, & Pigg	A waterlily, Rhizome sections were first identified as Ensete Seed masses first identified as .	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 has described from permineralized wood in the Princeton Chert.^[36] 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 representing the oldest confident Saururaceae species in the fossil record.^[37]

Family	Genus	Species	Authors	Notes	Images
Lauraceae	†	Undescribed^[18]		A lauraceous form species. Not described to species
Lauraceae	Ocotea	Undescribed^[18]		A stinkwood species. Not described to species
Lauraceae	Phoebe	Undescribed^[15]		A Phoebe species. Not described to species
Lauraceae	Sassafras	†Sassafras hesperia^[15]	Berry	A sassafras	Sassafras hesperia
Magnoliaceae	Magnolia	Undescribed^[38]		A magnolia, possibly Magnolia subgenus Talauma Not described to species
incertae sedis	†Dillhoffia	†Dillhoffia cachensis^[39]	Manchester & Pigg	A flower of uncertian 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 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 ,^[40] the alismatalean genus ,^[41] the alismatalean ,^[42] the asparagalean pollen morphogenus ,^[43] the lilialean genus ,^[44] and the poalean genus ,^[45]

Family	Genus	Species	Authors	Notes	Images
Amaryllidaceae	†	†^[46]	Pigg, , & DeVore	An onion relative
Araceae	Orontium	†Orontium wolfei^[47]	Bogner, Johnson, & Upchurch	A golden club	Orontium wolfei
Smilacaceae	Smilax	Undescribed^[18]		A greenbrier species. Not described to species.
Typhaceae	Typha	Undescribed^[18]		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.^[48] Benedict et al. (2011) described first fossils for the prunoid genus Oemleria along with the oldest Prunus flowers. The Prunus flowers are complimented by leaf fossils representeding five to six distinct morphotypes.^[49] Spiraea is known from an 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 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 had been described as of 2007.^[49] The rosaceous genus Physocarpus had been reported by Hopkins and Wehr (1994) as also occurring in the formation,^[38] 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.^[50]

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.^[51] 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.^[52] If fossils of the fruits and foliage in attachment are found, that would bring the species count down to three whole plant taxa.^[52] Additionally, the extinct genus 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.^[53] Wesley Wehr in 1994 reported Bignoniaceae seeds along with a single Rubiaceae fruit and an isolated Fabaceae leaf.^[54] 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.^[35]

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.^[55] That status was affirmed by Forest et al. (2005) who used both as fossil calibration points for phylogenetic analysis of Betulaceae.^[56]

Family	Genus	Species	Authors	Notes	Images
Anacardiaceae	Rhus	†Rhus boothillensis^[51]	Flynn, DeVore, & Pigg	A sumac, Hybridized with other Klondike Mountain Formation Rhus	Rhus boothillensis
Anacardiaceae	Rhus	†Rhus garwellii^[51]	Flynn, DeVore, & Pigg	A sumac, Hybridized with other Klondike Mountain Formation Rhus	Rhus garwellii
Anacardiaceae	Rhus	†Rhus malloryi^[15]^[51]	(Wolfe and Wehr) Flynn, DeVore & Pigg	A sumac, Hybridized with other Klondike Mountain Formation Rhus	Rhus malloryi
Anacardiaceae	Rhus	†Rhus republicensis^[51]	Flynn, DeVore, & Pigg	A sumac, Hybridized with other Klondike Mountain Formation Rhus
Aquifoliaceae	Ilex	Undescribed^[17]		A holly Not described to species
Araliaceae	Aralia	Undescribed^[23]		A spikenard species Not described to species.
Betulaceae	Alnus	^[15]	(Berry) Wolfe & Wehr	An Alder
Betulaceae	Betula	†Betula leopoldae^[15]^[57]	Wolfe & Wehr	A birch	Betula leopoldae
Betulaceae	Carpinus	†Carpinus perryae^[55]	Pigg, , & Wehr	A hornbeam
Betulaceae	Corylus	†Corylus johnsonii^[55]	Pigg, , & Wehr	A hazel nut	Corylus johnsonii
Betulaceae	†	†^[55]	Pigg, Manchester, & Wehr	A birch relative
Betulaceae	†	Undescribed^[55]		A birch relative different from the other Okanagan Highlands species	undescribed species
Bignoniaceae	Undescribed	Undescribed^[54]		A catalpa family member Not described to species	Bignoniaceae sp. fruit
Burseraceae	†Barghoornia	†Barghoornia oblongifolia^[15]	Wolfe & Wehr	An extinct Bursera relative	Barghoornia oblongifolia
Celtidaceae	Pteroceltis	Undescribed^[58]		A cannabaceous fruit Not described to species	Pteroceltis species fruit
Cercidiphyllaceae	Cercidiphyllum	†^[15]	(Dawson) Wolfe & Wehr	A katsura with suggested affinity to †,^[23] First described as "Populus" obtrita
Cornaceae	Cornus	Undescribed^[15]^[23]^[17]		A dogwood species, Not described to species
Elaeocarpaceae	Sloanea	Undescribed^[58]		An elaeocarpaceous fruit Not described to species	Sloanea sp.
Ericaceae	Arbutus	Undescribed^[38]^[17]		A madrone relative Not described to species
Ericaceae	Rhododendron	Undescribed^[38]^[23]^[17]		A rhododendron Not described to species
Ericaceae	Cf. Leucothoe	Undescribed^[38]		A doghobble relative Not described to species
Eucommiaceae	Eucommia	†Eucommia montana^[59]^[17]	Brown	A "hard rubber tree"	Eucommia montana
Fagaceae	†	Undescribed^[17]		A Castanea relative Not described to species
Fagaceae	†	†^[15]	(Knowlton) Hollick	A beech relative
Fagaceae	Fagus	†^[60]	Manchester & Dillhoff	A beech
Fagaceae	Quercus	Undescribed^[54]^[23]^[17]		An oak Not described to species
Grossulariaceae	Ribes	"Species 1"^[38]^[23]^[17]		A current Not described to species
Grossulariaceae	Ribes	"Species2"^[38]		A current Not described to species
Hamamelidaceae	Corylopsis	†Corylopsis reedae^[38]^[61]	, , & Wehr	A winter-hazel	Corylopsis reedae
Hamamelidaceae	Fothergilla	†Fothergilla malloryi^[61]	Radtke, Pigg, & Wehr	A witch alder	Fothergilla malloryi
Hamamelidaceae	†Langeria	†Langeria magnifica^[15]	Wolfe & Wehr	A witch hazel relative	Langeria magnifica
Hydrangeaceae	Hydrangea	Undescribed^[38]		A Hydrangea. Not described to species
Hydrangeaceae	Philadelphus	Undescribed^[38]		A mock-orange Not described to species
Icacinaceae		Unidentified^[54]^[23]		A Phytocrene relative Not described to species
Iteaceae	Itea	Undescribed^[15]^[17]		A virginia willow species Not described to species.
Juglandaceae	Carya	Undescribed^[23]^[17]		A walnut family relative. Not described to species.
Juglandaceae	†	†^[62]	(Brown) Manchester	A walnut family relative.
Juglandaceae	Juglans	Undescribed^[23]^[17]		A walnut family relative. Not described to species.
Juglandaceae	Pterocarya	Undescribed^[54]		A wing nut Not described to species
Lythraceae	Decodon	Undescribed^[23]		A swamp loosestrife Not described to species
Malvaceae	Craigia	Undescribed^[54]		A Craigia species Not described to species	Craigia sp.
Malvaceae	†Florissantia	†Florissantia quilchenensis^[63]	( & ) Manchester	A chocolate relative	Florissantia quilchenensis
Malvaceae	Hibiscus	Undescribed^[38]		A hibiscus Not described to species
Malvaceae	†	Undescribed^[54]		A relative Not described to species
Malvaceae	Tilia	†Tilia johnsoni^[15]	Wolfe & Wehr	A Linden	Tilia johnsoni
Malvaceae	Cf. Tilia	Undescribed^[58]		Linden relative fruits Not described to species	Cf. Tilia fruits
Menispermaceae	Calycocarpum	Undescribed^[35]		A moonseed Not described to species
Moraceae	Morus	Undescribed^[38]		A mulberry, two types known. Not described to species
Myricaceae	Comptonia	†Comptonia columbiana^[15]	Dawson	A Comptonia	Comptonia columbiana
Nyssaceae	†Tsukada	†Tsukada davidiifolia^[15]	Wolfe & Wehr	A dove-tree relative	Tsukada davidiifolia
Platanaceae	†	Undescribed^[54]	Manchester	A plane tree fruit taxon Not described to species	species
Platanaceae	†Macginitiea	†Macginitiea gracilis^[15]	(Lesquereux) Wolfe & Wehr	A plane tree relative	Macginitiea gracilis
Platanaceae		Undescribed^[58]		A sycamore stamen head isolated stamen clusters placed as Not described to species	stamen cluster
Polygalaceae		Undescribed^[54]		A milkwort relative Not described to species
Ranunculaceae	Clematis	Undescribed^[38]		A Clematis Not described to species	Clematis species
Rosaceae	Amelanchier	Undescribed^[38]		A service berry Not described to species
Rosaceae	Malus	Undescribed^[17]		An apple Not described to species
Rosaceae	Neviusia	Undescribed^[38]		A snow-wreath Not described to species
Rosaceae	Oemleria	†^[48]	, DeVore, & Pigg	An Osoberry
Rosaceae	Photinia	†^[15]	Wolfe & Wehr	A Christmas-berry relative
Rosaceae	Prunus	†^[48]	Benedict, DeVore, & Pigg	A cherry relative
Rosaceae	Prunus	"Species 1"^[15]		A prunoid leaf Not described to species
Rosaceae	Prunus	"Species 2"^[15]		A prunoid leaf Not described to species
Rosaceae	Prunus	"Species 3"^[15]		A prunoid leaf Not described to species
Rosaceae	Pyracantha	Undescribed^[49]		A firethorn sp.^[49] Tentative record, Not described to species.
Rosaceae	Rubus	Undescribed^[38]		A blackberry Not described to species
Rosaceae	Spiraea	Undescribed^[23]^[17]		A bridal wreath Not described to species.
Rosaceae	Aff. Sorbus	Undescribed^[23]^[17]		A rowan relative Not described to species.
Rosaceae	Aff. Physocarpus	Undescribed^[23]		A possible nine-bark Not described to species Possibly stem Neillieae^[50]
Rosaceae	Cf. Crataegus	Undescribed^[38]		A hawthorn relative Not described to species
Rosaceae	Cf. Malus	Undescribed^[49]		A maloid species possibly apple or pear Not described to species
Rosaceae	Cf. Spiraea	Undescribed^[49]		A bridal wreath relative Not described to species
Rosaceae	Cf. Pyrus	Undescribed^[49]		A maloid species possibly apple or pear Not described to species
Sabiaceae	Meliosma	Undescribed^[35]		A Meliosma species Not described to species
Sabiaceae		Undescribed^[35]		A species Not described to species
Salicaceae	Populus	Undescribed^[23]^[17]		A cottonwood Not described to species First identified as †^[7]
Salicaceae		Undescribed^[23]		A Not described to species
Salicaceae	Salix	Undescribed^[23]^[17]		A willow Not described to species
Sapindaceae	Acer?	^[64]	Heer, 1876	A possible maple
Sapindaceae	Acer	Acer hillsi^[64]	Wolfe &	A maple
Sapindaceae	Acer	Acer republicense^[64]	Wolfe & Tanai	A maple
Sapindaceae	Acer	Acer spitzi^[64]	Wolfe & Tanai	A maple
Sapindaceae	Acer	Acer stonebergae^[64]	Wolfe & Tanai	A maple
Sapindaceae	Acer	Acer toradense^[64]	Wolfe & Tanai	A maple
Sapindaceae	Acer	Acer washingtonense^[64]	Wolfe & Tanai	A maple
Sapindaceae	Acer	^[64]	Wolfe & Tanai	A maple
Sapindaceae	Aesculus	Undescribed^[23]^[65]		A horse chestnut Not described to species.
Sapindaceae	†	†^[15]	(Brown) Wolf & Wehr	An extinct sapindalean species
Sapindaceae	Dipteronia	†Dipteronia brownii^[66]	&	A Dipteronia	Dipteronia brownii
Sapindaceae	Koelreuteria	†^[67]	Wang et al.	A Koelreuteria species
Sapindaceae	Cf. Boniodendron	†^[15]^[67]	Becker	A sapindaceous species first described as a Koelreuteria species, considered Cf. Boniodendron by Wang et al (2012).
Schisandraceae	Kadsura	Undescribed^[18]		A kadsura species. Not described to species
Schoepfiaceae	Schoepfia	†^[15]	(LaMotte) Wolfe & Wehr	A possible Schoepfia relative, First described as "Cornus acuminata.
Theaceae		"Species A"^[15]		A theaceous species similar to Gordonia Not described to species
Theaceae		"Species B"^[15]		A theaceous species similar to Cleyera Not described to species
Trochodendraceae?	†	Undescribed^[54]		A trochodendroid of uncertain placement.^[53] Fruits of the leaf taxon Not described to species.
Trochodendraceae	†Paraconcavistylon	†Paraconcavistylon wehrii^[52]^[53]	( et al.)	A Trochodendrale first described as "Concavistylon" wehrii moved to a new genus in 2020.
Trochodendraceae	†Pentacentron	†Pentacentron sternhartae^[52]	et al.	A Trochodendrale	Pentacentron sternhartae
Trochodendraceae	Tetracentron	†Tetracentron hopkinsii^[52]	Pigg et al.	A Trochodendrale, possibly the leaves of Pentacentron sternhartae	Tetracentron hopkinsii
Trochodendraceae	Trochodendron	†Trochodendron nastae^[68]	Pigg, Wehr, &	A Trochodendron Possibly the leaves of Trochodendron drachukii	Trochodendron nastae
Trochodendraceae	†	Undescribed^[23]		A trochodendroid of uncertain placement.^[53] Leaves of the fruit taxon Not described to species.
Ulmaceae		Undescribed^[35]		An elm relative Not described to species
Ulmaceae	Ulmus	†^[69]	(Berry) LaMotte	An elm species
Ulmaceae	Ulmus	†Ulmus okanaganensis	Denk & Dillhoff	An elm species, the fruits were first identified as Ulmus section Chaetoptelea.	Ulmus okanaganensis
Vitaceae	Vitis	Undesribed^[38]^[54]		Grape seeds^[54] and leaves^[38] Not described to species
Incertae sedis	†	Undescribed^[23]		A leaf morphotype of uncertain affiliation. Not described to species.
incertae sedis	†	†^[54]	Crane	A winged fruit of unidentified affinities
incertae sedis	†Pteronepelys	†Pteronepelys wehrii^[70]	Manchester	A samara of uncertain affinities.	Pteronepelys wehrii
†incertae sedis		†^[15]	Wolfe & Wehr	An incertae sedis angiosperm possibly of Hamamelididae affiliations

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.^[71] The most prevalent orders are Diptera and Hemiptera, each making up approximately 30% of the fossil insects known in 1992.

Blattodea[]

Family	Genus	species	Notes	Images
Blaberidae	Undescribed	Undescribed^[17]	A cockroach Not described to genus/species
Blattoidae	Undescribed	Undescribed^[71]	A blattoidean cockroach Not described to genus/species	Undescribed Blattoidea
Isoptera	Undescribed	Undescribed^[71]	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,^[71] but the beetle fauna has not been described in depth yet, with only two weevil species having been fully described.^[72]^[73] A third beetle group belonging to the bean beetle tribe has been identified as palm beetles of the – genus group.^[74]

Family	Genus	Species	Authors	Notes	Images
Brentidae	†	†^[72]	Legalov, 2013	An Ithycerinae weevil
Brentidae	†	†^[73]	Legalov, 2015	An Ithycerinae weevil
Carabidae	Undescribed	Undescribed^[71]		A ground beetle Not described to genus/species	Unidentified Carabidae
Cerambycidae	Undescribed	Undescribed^[71]		A long-horn beetle Not described to genus/species
Chrysomelidae	– genus group	Undescribed^[74]		palm beetles in the tribe . Not described to genus/species
Dytiscidae	Undescribed	Undescribed^[71]		A diving beetle Not described to genus/species
Elateridae	Undescribed	Undescribed^[71]		A click beetle Not described to genus/species
Lucanidae	Undescribed	Undescribed^[71]		A stag beetle Not described to genus/species

Dermaptera[]

The order Dermaptera was first reported in 1992^[71] 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.^[75]

Family	Genus	Species	Authors	Notes	Images
Forficulidae?	Unidentified	"Forficulid species 1"^[75]		A possible forficulid earwig with long cerci Not described to genus/species	"Forficulid species 1"
Forficulidae?	Unidentified	"Forficulid species 2"^[75]		A possible forficulid earwig with short cerci Not described to genus/species

Diptera[]

Family	Genus	Species	Authors	Notes	Images
Bibionidae	Undescribed	Undescribed^[71]		A march fly Not described to genus/species	unidentified Bibionidae
Mycetophilidae	Undescribed	Undescribed^[71]		A fungus gnat Not described to genus/species
Pipunculidae	†Metanephrocerus	†Metanephrocerus belgardeae^[76]	Archibald, , & Mathewes, 2014	A pipunculid big-headed fly	Metanephrocerus belgardeae
Tipulidae	Undescribed	Undescribed^[71]		A crane fly Not described to genus/species	Undescribed Tipulidae

Ephemeroptera[]

Lewis (1992) listed one species of Heptageniidae mayflies and three specimens that he did not place to family.^[71] The Heptageniidae specimen was later described as a squaregill mayfly by Sinitchenkova (1999) and the oldest member of the genus Neoephemera.^[77]

Family	Genus	Species	Authors	Notes	Images
Neoephemeridae	Neoephemera	†Neoephemera antiqua^[77]	, 1999	A squaregill mayfly, first tentatively identified as a flat headed mayfly.^[71]	Neoephemera antiqua

Hemiptera[]

Family	Genus	Species	Notes	Images
Aphididae	Undescribed	Undescribed^[71]	An aphid Not described to genus/species.
Aphrophoridae	Aphrophora	Undescribed^[71]	An aphrophorid spittlebug Not described to species.	Aphrophora species
Aphrophoridae	†	Undescribed^[71]	An aphrophorid spittlebug Not described to species.
Cercopidae	†	Undescribed^[71]	A cercopid froghopper Not described to species.
Cercopidae	Undescribed	Undescribed^[71]	A cercopid froghopper Not described to genus/species.	undescribed Cercopidae
Fulgoroidea	Undescribed	Undescribed^[71]	A frog hopper Not described to genus/species.
Pentatomidae	Undescribed	Undescribed^[71]	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.^[78]

Family	Genus	Species	Authors	Notes	Images
Angarosphecidae	†	Undescribed^[78]		An angarosphecid spheciform wasp. Not described to species
Angarosphecidae	Undescribed	Undescribed^[78]		An angarosphecid spheciform wasp Likely not . Not described to species
Braconidae	Undescribed	Undescribed^[71]^[78]		braconid parasitic wasps Not described to genus/species.	Unidentified Braconidae
Cimbicidae	Undescribed	Undescribed^[78]		A cimbicid sawfly. Not Described to genus/species	Unidentified Cimbicidae
Diapriidae	Undescribed	Undescribed^[78]		A diapriid diaprioid wasp Not described to species
Formicidae	†Klondikia	†Klondikia whiteae^[79]	Dlussky & Rasnitsyn, 2003	An ant of uncertain subfamily affiliation
Formicidae	Myrmeciites	"Indesterminate"^[80]	Archibald, , & , 2006	A bulldog ant form genus	sp.
Formicidae	Oecophylla	†^[81]	(Dlussky & Rasnitsyn, 1999)	An ant, described as Camponotites kraussei, Moved to Oecophylla kraussei in 2017^[82]
Formicidae	†Propalosoma	†Propalosoma gutierrezae^[81]	Dlussky & Rasnitsyn, 1999	A bulldog ant, first described as a Rhopalosomatidae wasp, moved to myrmeciinae in 2018^[83]	Propalosoma gutierrezae
Formicidae	Undescribed	Undescribed^[78]		Ants of uncertain subfamily placement.^[78]	Unidentified formicidae
Halictidae (?)	Undescribed	Undescribed^[78]		A possible sweat bee Not described to genus/species
Ichneumonidae	Undescribed	Undescribed^[78]		ichneumonid parasitic wasps unplaced to subfamily Not described to genus/species	Undescribed Ichneumonidae
Megachilidae	Undescribed	Undescribed^[78]		Megachilid leaf-cutter bee herbivory trace fossils on leaves Not described to genus/species
Pamphiliidae	†Ulteramus	†Ulteramus republicensis^[84]	Archibald & Rasnitsyn, 2015	A parasitic wasp
Pompilidae	Undescribed	Undescribed^[78]		A pompilid spider wasp Not described to genus/species
Proctotrupidae	Undescribed	Undescribed^[78]		A proctotrupid parasitic wasp Not described to genus/species
Roproniidae	Undescribed	Undescribed^[78]		A roproniid (sensu lato) proctotrupoid wasp Not described to genus/species
Scoliidae	Undescribed	Undescribed^[78]		An scoliid wasps Not described to genus/species
Sphecidae	Undescribed	Undescribed^[78]		A sphecid (sensu stricto) wasp Not described to genus/species
Tenthredinidae	Undescribed	Undescribed^[78]		A Tenthredinid sawfly Not described to genus/species
Vespidae	Undescribed	Undescribed^[78]		A vespid wasp Not described to genus/species

Lepidoptera[]

A solitary lepidopteran 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]

Family	Genus	Species	Authors	Year	Notes	Images
Erebidae	Undescribed	Undescribed^[85]		1986	An arctiine tiger moth Not described

Mecoptera[]

A number of mecopteran species belonging to the families Cimbrophlebiidae, Dinopanorpidae, Eorpidae, and Panorpidae are also known.^[86]

Family	Genus	Species	Authors	Year	Notes	Images
†Cimbrophlebiidae	†Cimbrophlebia	†^[86]	Archibald, 2009	2009	A Cimbrophlebiid scorpionfly
†Cimbrophlebiidae	†Cimbrophlebia	†^[86]	Archibald, 2009	2009	A Cimbrophlebiid scorpionfly
†Dinopanorpidae	†Dinokanaga	†Dinokanaga andersoni^[87]	Archibald, 2005	2005	A scorpion fly species	Dinokanaga andersoni
†Dinopanorpidae	†Dinokanaga	†Dinokanaga dowsonae^[87]	Archibald, 2005	2005	A scorpion fly species
†Dinopanorpidae	†Dinokanaga	†Dinokanaga sternbergi^[87]	Archibald, 2005	2005	A scorpion fly species
†Eorpidae	†Eorpa	†Eorpa elverumi^[88]	Archibald, Mathewes, & Greenwood, 2013	2013	A mecopteran scorpionfly	Eorpa elverumi
†Eorpidae	†Eorpa	?†Eorpa ypsipeda^[88]	Archibald, Mathewes, & Greenwood, 2013	2013	A mecopteran scorpionfly, tentatively identified	Possible E. ypsipeda^[88]

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.^[5]

Family	Genus	Species	Authors	Year	Notes	Images
Chrysopidae	†	†^[89]	& , 2013	2013	A green lacewing
Hemerobiidae	†Proneuronema	†Proneuronema wehri^[90]^[91]	(Makarkin, Archibald, & , 2003)	2003	A hemerobiid lacewing, originally placed in †Cretomerobius, moved to †Proneuronema in
Ithonidae	†Allorapisma	†Allorapisma chuorum^[92]	Makarkin & Archibald, 2009	2009	A moth lacewing	Allorapisma chuorum
Ithonidae	†Palaeopsychops	†Palaeopsychops marringerae^[93]	Archibald & Makarkin, 2006	2006	A polystechotid group^[94] moth lacewing	Palaeopsychops marringerae
Ithonidae	†Palaeopsychops	†Palaeopsychops timmi^[93]	Archibald & Makarkin, 2006	2006	A polystechotid group^[94] moth lacewing	Palaeopsychops timmi
Ithonidae	†	†^[93]	Archibald & Makarkin, 2006	2006	A polystechotid group^[94] moth lacewing
Ithonidae	†	†^[93]	Archibald & Makarkin, 2006	2006	A polystechotid group^[94] moth lacewing
Ithonidae	†	†^[93]	Archibald & Makarkin, 2006	2006	A polystechotid group^[94] moth lacewing
Nymphidae	?Nymphes	†Nymphes? georgei^[95]	Archibald, Makarkin, & Ansorge, 2009	2009	A nymphid lacewing, possibly a species of †^[96]	Nymphes georgei
Osmylidae	†	†^[97]	Makarkin, Archibald, & Mathewes, 2021	2021	A osmylid lacewing
Psychopsidae?	†Ainigmapsychops	†Ainigmapsychops inexspectatus^[5]	Makarkin & Archibald, 2014	2014	A possible psychopsid lacewing	Ainigmapsychops inexspectatus

Odonata[]

Family	Genus	Species	Authors	Year	Notes
Aeshnidae	†	†^[98]	Archibald & Cannings, 2019	2019	A darner dragonfly
Aeshnidae	†	†^[98]	Archibald & Cannings, 2019	2019	A darner dragonfly
Aeshnidae	†	†^[98]	Archibald & Cannings, 2019	2019	A darner dragonfly
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
†	†	†^[99]	Archibald & Cannings, 2021	2021	A odonate
Euphaeidae	†	†^[100]	Archibald & Cannings, 2021	2021	A gossamerwing damselfly. Not to be confused with the plant , also from the formation
†	†	†^[99]	Archibald & Cannings	2021	A possibly odonate

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.^[101] 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.^[102]

Family	Genus	Species	Authors	Year	Notes	Images
†Susumaniidae	†Eoprephasma	†Eoprephasma hichensi^[101]	Archibald & Bradler, 2015	2015	A Susumaniinae stick insect species

Raphidioptera[]

Family	Genus	Species	Authors	Year	Notes	Images
Raphidiidae	†	†^[103]	Archibald & Makarkin, 2021	2021	A raphidiid snakefly

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.^[104] 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, , 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.^[105] 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 quadrangle, Ferry and Okanogan Counties, Washington. The fossils were tentatively identified by paleoichthyologist David Dunkle as members of the genera Amyzon, , and an undefined salmonid.^[106] 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.^[107] 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.^[108]^[109] 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.^[106] A year later the percopsid fossils were also described as Libotonius pearsoni, extending the range of the genus south from the Allenby Formation.^[108]

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	, 1977	1977	A bowfin, known from isolated scales	"Amia" hesperia scale
Catostomidae	†Amyzon	†Amyzon aggregatum	Wilson, 1977	1977	A sucker
Catostomidae	†Amyzon	Unidentified^[110]	(Wilson, 1977)	1977	A sucker, originally identified as Amyzon aggregatum
Salmonidae	†Eosalmo	†Eosalmo driftwoodensis^[111]	Wilson, 1977	1999	A Salmon	Eosalmo driftwoodensis
Hiodontidae	Hiodon	†Hiodon woodruffi^[106]	Wilson, 1978	1978	A mooneye, first described as "Eohiodon" woodruffi.^[106]	Hiodon woodruffi
†Libotoniidae	†Libotonius	†Libotonius pearsoni^[108]	Wilson, 1979	1979	A sand roller relative.	Libotonius pearsoni
incertae sedis (Aves)	Unidentified	"Unnamed"^[112]		2019	indeterminate feathers and a skeleton	Unidentified feather

References[]

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