Allenby Formation

Allenby Formation
Allenby Formation
Stratigraphic range: Ypresian; 52.5–48 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Alternating Princeton Chert and coal sequences
Type	Geological formation
Unit of	Princeton Group, Eocene Okanagan Highlands
Sub-units	Princeton Chert, Vermillion Bluffs Shale
Overlies	Cedar Formation
Area	300 km2 (120 sq mi)
Thickness	1,860–2,100 m (6,100–6,890 ft)
Lithology
Primary	Shale, sandstone
Other	Coal–breccia, coal–chert
Location
Coordinates	49°22.6′N 120°32.8′W / 49.3767°N 120.5467°W
Approximate paleocoordinates	53°06′N 107°30′W / 53.1°N 107.5°W
Region	British Columbia
Country	Canada
Extent	Princeton Basin & Tulameen basin
Type section
Named for	Allenby, British Columbia
Named by	Shaw
Year defined	1952
	Allenby Formation (Canada) Allenby Formation (British Columbia)

The Allenby formation is a sedimentary rock formation in British Columbia which was deposited during the Ypresian stage of the Early Eocene. It consists of conglomerates, sandstones with interbedded shales and coal. The shales contain an abundance of insect, fish and plant fossils known from 1877 and onward, while the Princeton Chert was first indented in the 1950s and is known from anatomically preserved plants.

There are several notable fossil producing localities in the Princeton & Tulameen basins. Historical collection sites included Nine Mile Creek, Vermilian Bluffs, and Whipsaw Creek, while modern sites include One Mile Creek, Pleasant Valley, Thomas Ranch, and the Princeton Chert.

Extent and correlation

The Allenby is estimated to have an overall extent of approximately 300 km² (120 sq mi), though actual outcroppings of the formation make up less than 1% of the formation, while other exploratory contact is via boreholes and mines. The half-graben which contains the formation is separated into two major depositional basins, the Princeton basin around Princeton, British Columbia and the Tulameen basin centered approximately 17 km (11 mi) west. The grabens extensional faults at the eastern side of the basin place the hanging wall Allenby strata in contact with much older foot wall strata of the Nicola Formation which dates to the Upper Triassic.^[1]^[2]^[3]

The Allenby Formation is the southern-most of the Eocene Okanagan Highlands lakes in British Columbia, and second most southern site after the Klondike Mountain Formation of Republic, Washington and northern Ferry County. In British Columbia, the formation is coeval to the Tranquille Formation, known from the McAbee Fossil Beds and Falkland site, the Coldwater Beds, known from the Quilchena site, and Driftwood Canyon Provincial Park. The highlands, including the Allenby Formation, have been described as one of the "Great Canadian Lagerstätten"^[4] based on the diversity, quality and unique nature of the biotas that are preserved. The highlands temperate biome preserved across a large transect of lakes recorded many of the earliest appearances of modern genera, while also documenting the last stands of ancient lines.^[4]

The warm temperate uplands floras of the Allenby Formation and the highlands, associated with downfaulted lacustrine basins and active volcanism are noted to have no exact modern equivalents, due to the more seasonally equitable conditions of the Early Eocene. However, the formation has been compared to the upland ecological islands in the Virunga Mountains within the Albertine Rift of the African rift valley.^[5]

The earliest work in the region was on exploratory expeditions in 1877 and 1878, with fossils collected in the areas of Nine-Mile Creek, Vermilian Bluffs on the Similkameen River, and Whipsaw Creek. While reporting on additional plant fossils collected from British Columbia, Penhallow (1906) noted the likely coeval status of the Princeton basins with many of the sites now considered the Okanagan Highlands.^[6] Modern collecting has centered on the areas around One Mile Creek, Pleasant Valley, and Thomas Ranch.^[2]

Age

The age estimates for the Allenby Formation have varied a number of times since the first explorations happened in the 1870s. Shaw (1952) dated the formation as Oligocene, an age followed by Arnold (1955).^[7]^[8] Half a decade later, the older age of 48 ± 2 million years old was first suggested, with a younger age being suggested at 46.2 ± 1.9 million years old in 2000 and an older date of 52.08 ± 0.12 million years ago obtained from uranium–lead dating of zircons from Vermilion Bluffs shale in 2005.^[1]

Lithology

The Allenby is composed of cyclical sedimentation events that were deposited along the course of a river-system in conjunction with depositional areas from nearby lakes and wetlands. Coeval volcanic eruptive events are recorded as interbeds of tephras and lavas, while the riverine course is marked with depositional areas of conglomerates and sandstones. The quieter environments are noted for finer layers of shales and coalified layers.^[1]

The coal seams throughout the formation are typically sub-bituminous.^[1]

Notable in conjunction with the coal seams are sections of chert which formed during silica rich periods. The rapid cyclical changes from coal to chert and back are not noted in any other fossil locality in the world. An estimated 49 coal-chert cycles are known, though the exact conditions for this process are not well understood. Silica rich volcanic episodes in the region during deposition would have been needed for formation of the cherts, while slowly moving waters and gently subsiding terrains would be needed for the peats and fens to accumulate. Rates of organic deposition in swamps have been estimated at 0.5–1 mm (0.020–0.039 in) in modern temperate climates, this suggests the time needed for each 10–20 cm (3.9–7.9 in) chert layer would be at least 100 years or more, with the full sequence of cycles taking place over no more than 15,000 years.^[1]

Palynoflora

Palynological analysis of samples from the Thomas ranch site by Dillhoff et al. (2013) resulted in the identification of 32 pollen and spore types that were assignable to family or genus level, with a total number of distinct pollen and spore types, including unassignable morphotypes, number over 70. The predominant pollens of the site are conifers, which make up between 85%–97% of the total pollens, while the angiosperm pollens are dominated by members of Betulaceae.^[2]

Several pteridophyte families and genera are represented as spore fossils alone, without corresponding megafossil records, including Lycopodiaceae, Osmundaceae, and Schizaeaceae. Similarly, at least three additional conifer genera are only present as pollen fossils and up to 12 angiosperms are present in the pollen record. Sometimes considered a Biostratgraphic index fossil, the angiosperm palynospecies Pistillipollenites macgregorii has been recovered from several sites in the Allenby Formation, while the palynospecies Erdtmanipollis pachysandroides is rare, having only been reported from the formation twice.^[2]

Family	Genus	Species	Pollen/Macrofossil	Notes
Aquifoliaceae	Ilex	Unidentified^[2]	Pollen	A holly palynomorph
Arecaceae	Sabal	Cf. †Sabal granopollenites^[2]	Pollen	A palm palynomorph
Buxaceae	†Erdtmanipollis	†Erdtmanipollis pachysandroides^[2]	Pollen	A box family palynomorph
Betulaceae	Alnus	Unidentified^[2]	Pollen & macrofossils	An alder palynomorph
Betulaceae	Betula	Unidentified^[2]	Pollen & macrofossils	A birch palynomorph
Betulaceae	Carpinus	Unidentified^[2]	Pollen	A hornbeam palynomorph
Betulaceae	Corylus	unidentified^[2]	Pollen	A hazelnut palynomorph
Cupressaceae	Cunninghamia	unidentified^[2]	Pollen	A Cunninghamia like palynomorph
Cupressaceae	Sequoiapollenites	Unidentified^[2]	Pollen & macrofossils	A redwood palynomorph
Cupressaceae	Taxodiaceaepollenites	Unidentified^[2]	Pollen & macrofossils	A Taxodioideae subfamily palynomorph
Elaeagnaceae	Cf. Elaeagnus	Unidentified^[2]	Pollen	An elaeagnaceous palynomorph, similar to oleaster
Ericaceae	unidentified	Unidentified^[2]	Pollen	An ericaceous palynomorph of uncertain affinity
Fagaceae	Castanea	Unidentified^[2]	Pollen	A chestnut palynomorph
Fagaceae	†Eotrigonobalanus	Unidentified^[9]	Pollen	A fagaceous palynomorph
Fagaceae	Fagus	"Fagus Pollen type 3"^[9]	Pollen	A beech palynomorph
Fagaceae	Fagus	"Fagus Pollen type 2"^[9]	Pollen	A beech palynomorph
Fagaceae	†Paraquercus	†Paraquercus eocaena^[9]	Pollen	A fagaceous palynomorph
Fagaceae	Quercus	"Quercus Pollen type 1"^[9]	Pollen	An oak palynomorph, similar to Quercus Group Lobatae pollen
Fagaceae	Quercus	"Quercus Pollen type 2"^[9]	Pollen	An oak palynomorph, ancestral type with Quercus Group Ilex morphology
Fagaceae	†Trigonobalanopsis	Unidentified^[9]	Pollen	A fagaceous palynomorph
Fagaceae	Unidentified	Unidentified^[9]	Pollen	A Fagoideceous palynomorph
Fagaceae	Cf. Quercus	Unidentified^[2]	Pollen	A fagaceous palynomorph, similar to oak
Ginkgoaceae	†Cycadopites	†Cycadopites follicularis^[2]	Pollen & macrofossils	A Gingko palynomorph
Hamamelidaceae	Liquidambar	Unidentified^[2]	Pollen	A sweet gum palynomorph
Juglandaceae	Carya	Unidentified^[2]	Pollen	A hickory palynomorph
Juglandaceae	Pterocarya	Unidentified^[2]	Pollen	A hickory palynomorph
Lycopodiaceae	Lycopodium	Unidentified^[2]	Pollen	A lycopod palynomorph
Malvaceae	Tilia	Unidentified^[2]	Pollen	A linden palynomorph
Osmundaceae	Osmundasporites	Unidentified^[2]	Pollen	An osmundaceous fern palynomorph
Pinaceae	Abies	Unidentified^[2]	Pollen & macrofossils	A fir palynomorph
Pinaceae	†Alisporites	Unidentified^[2]	Pollen	A pine family palynomorph
Pinaceae	Picea	Unidentified^[2]	Pollen	A Picea palynomorph
Pinaceae	Pinus	Unidentified^[2]	Pollen & macrofossils	A Pinus palynomorph
Pinaceae	†Pityosporites	Unidentified^[2]	Pollen	A pine family palynomorph
Pinaceae	Pseudolarix	Unidentified^[2]	Pollen & macrofossils	A Pseudolarix palynomorph
Pinaceae	Tsuga	Unidentified^[2]	Pollen	A Tsuga palynomorph
Pinaceae	Cf. Larix	Unidentified^[2]	Pollen	A Laricoidae palynomorph, similar to larch
Pinaceae	Cf. Pseudotsuga	Unidentified^[2]	Pollen	A Laricoidae palynomorph, similar to pseudotsuga
Platanaceae	Platanus	Unidentified^[2]	Pollen	A Platanus palynomorph
Potamogetonaceae	Potamogeton	Unidentified^[2]	Pollen	A Potamogeton palynomorph
Rosaceae	Unidentified	Unidentified^[2]	Pollen & macrofossils	Rose famnily palynomorphs
Salicaceae	Salix	Unidentified^[2]	Pollen	A willow palynomorph
Salviniaceae	Azolla	Unidentified^[2]	Pollen & macrofossils	A duck weed palynomorph
Sapindaceae	Acer	Unidentified^[2]	Pollen & macrofossils	A maple palynomorph
Sapindaceae	Aesculus	Unidentified^[2]	Pollen & macrofossils	A horse chestnut palynomorph
Sapotaceae	Unidentified	Unidentified^[2]	Pollen	A sapotaceous palynomorph
Taxaceae	Taxus	Unidentified^[2]	Pollen	A yew palynomorph
Ulmaceae	Ulmus	Unidentified^[2]	Pollen & macrofossils	An elm palynomorph
incertae sedis	†Pistillipollenites	†Pistillipollenites macgregorii^[2]	Pollen	A palynomorph of uncertain affinity, possibly a Gentianaceae or Euphorbiaceae species

Compression paleobiota

A group of six mosses were described from the Allenby Formation by Kuc (1972, 1974) representing the genera Ditrichites, Hypnites and Plagiopodopsis, with two species placed in the morphogenus Muscites.^[10]^[11] Dillhoff et al. (2013) identified twelve distinct gymnosperm taxa spanning the families Cupressaceae, Ginkgoaceae, and Pinaceae. While being the minority component of the Thomas Ranch flora by total fossil numbers, angiosperms have a higher diversity, with 45 distinct morphotypes represented as foliage, reproductive structures, or both. Seventeen of the morphotypes are identifiable to genus or species, with members of the family Betulaceae being most prominent. At least common one leaf type is suggested to possibly represent an extinct plant order, but has not been described.^[2] Only two pteridophyte species have been described from the compression flora, Azolla primaeva by Penhallow (1890) and Equisetum similkamense by Dawson (1878).^[12]^[8]

The following fossil conifers, pteridophytes, ginkgophytes and bryophytes have been described from the Allenby Formation:

Bryophytes

Family	Genus	Species	Authority	Notes
Amblystegiaceae	Hypnites	†Hypnites jovet-astiae^[11]^[13]	(Kuc) Miller	An amblystegiaceous moss First described as Palaeohypnum jovet-asti, moved to Hypnites jovet-astiae in 1980
Amblystegiaceae	Hypnites	†Hypnites steerei^[11]^[13]	(Kuc) Miller	An amblystegiaceous moss First described as Palaeohypnum steerei, moved to Hypnites steerei in 1980
Bartramiaceae	†Plagiopodopsis	†Plagiopodopsis eocenicus^[10]^[13]	(Kuc) Miller	A bartramiaceous moss First described as Muscites eocenicus, moved to Plagiopodopsis eocenicus in 1980
?Ditrichaceae	†Ditrichites	†Ditrichites fylesi^[11]	Kuc	A Ditrichaceous moss
incertae sedis	†Muscites	†Muscites maycocki^[11]	Kuc	A moss of uncertain placement
incertae sedis	†Muscites	†Muscites ritchiei^[11]	Kuc	A moss of uncertain placement

Pteridophytes

Family	Genus	Species	Authority	Notes	Images
Equisetaceae	Equisetum	†Equisetum similkamense^[12]	Dawson	A scouring rush
Salviniaceae	Azolla	†Azolla primaeva^[8]	(Penhallow) Arnold	A mosquito fern First described as Azollophyllum primaevum

Gingkophytes

Family	Genus	Species	Authority	Notes	Images
Ginkgoaceae	Ginkgo	Ginkgo biloba^[14]^[2]	Linnaeus	A ginkgo
Ginkgoaceae	Ginkgo	†Ginkgo dissecta^[14]^[2]	Mustoe, 2002	A ginkgo with highly dissected leaves

Pinophytes

Family	Genus	Species	Authority	Notes	Images
Cupressaceae	Chamaecyparis	†Chamaecyparis linguaefolia^[15]	(Lesquereux) MacGinitie	A false cypress
Cupressaceae	Metasequoia	†Metasequoia occidentalis^[16]^[2]	(Newberry) Chaney	A dawn redwood First identified as "Sequoia" brevifolia, "S." heeri. "S." langsdorfii (in part), "S." nordenskiöldi, & Taxodium distichum miocenum (in part)
Cupressaceae	Sequoia	†Sequoia affinis^[15]	Lesquereux	A redwood	Sequoia affinis
Cupressaceae	Taxodium	†Taxodium dubium^[16]	(Sternberg) Heer	A bald cypress First identified as "Sequoia" angustifolia, "S." langsdorfii (in part), & Taxodium distichum miocenum (in part)
Pinaceae	Abies	†Abies milleri^[2]	Shorn & Wehr, 1986	Oldest true fir described	Abies milleri
Pinaceae	Picea	Undescribed^[15]	Miller	A spruce Not described to species
Pinaceae	Pinus	†Pinus latahensis^[15]	Berry	A 5-needle pine	Pinus latahensis
Pinaceae	Pinus	†Pinus monticolensis^[15]	Berry	A pinaceous winged seed
Pinaceae	Pinus	†Pinus trunculus^[15]	Dawson, 1890	A 3-needle pine
Pinaceae	Pinus	†Pinus tulameenensis^[15]	Penhallow	A 5-needle pine
Pinaceae	Pseudolarix	Pseudolarix amabilis^[17]	(J.Nelson) Rehder	A golden larch Originally identified as Pseudolarix americana,^[15] then as Pseudolarix arnoldii^[18]
Pinaceae	Pseudolarix	†Pseudolarix wehrii^[17]	Gooch	A golden larch	Pseudolarix wehrii

Angiosperms

Family	Genus	Species	Authority	Notes	Images
Anacardiaceae	Rhus	†Rhus malloryi^[19]	Wolfe & Wehr	A sumac	Rhus malloryi
Amaryllidaceae	†Paleoallium	†Paleoallium billgenseli^[20]	Pigg, Bryan, & DeVore	An onion relative	Paleoallium billgenseli
Araceae	Orontium	†Orontium wolfei^[21]	Bogner, Johnson, Kvaček & Upchurch	A golden-club	Orontium wolfei
Betulaceae	Alnus	†Alnus parvifolia^[19]^[2]	(Berry) Wolfe & Wehr	An Alder	Alnus parvifolia
Betulaceae	Betula	†Betula leopoldae^[19]^[2]	Wolfe & Wehr	A birch	Betula leopoldae
Betulaceae	†Palaeocarpinus	†Palaeocarpinus stonebergae^[22]	Pigg, Manchester, & Wehr	A coryloid genus
Cercidiphyllaceae	Cercidiphyllum	†Cercidiphyllum obtritum^[19]	(Dawson) Wolfe & Wehr	A katsura	Cercidiphyllum obtritum
Fagaceae	†Fagopsis	†Fagopsis undulata^[19]	(Knowlton) Wolfe & Wehr	A beech	Fagopsis undulata
Fagaceae	Fagus	Undescribed^[2]		A beech species Not described to species
Grossulariaceae	Ribes	Undescribed^[2]		A gooseberry species Not described
Hamamelidaceae	Fothergilla	†Fothergilla malloryi^[2]	Radtke, Pigg & Wehr	A winter-hazel species	Fothergilla malloryi
Juglandaceae	Pterocarya	Undescribed^[23]		A wingnut Not described to species
Lauraceae	Lindera	Undescribed^[23]		A laural species Not described to species
Lauraceae	Sassafras	†Sassafras hesperia^[19]^[2]	Berry	A laural species	Sassafras hesperia
Malvaceae	Florissantia	Undescribed^[24]		An extinct sterculioid flower Not described to species
Myricaceae	Comptonia	†Comptonia columbiana^[12]^[2]	Dawson	A sweet fern	Comptonia columbiana
Nyssaceae	†Tsukada	†Tsukada davidiifolia^[19]^[2]	Wolfe & Wehr	A dove-tree relative	Tsukada davidiifolia
Platanaceae	Macginicarpa	Undescribed^[2]	Manchester	A sycamore morphospecies
Platanaceae	†Macginitiea	†Macginitiea gracilis^[19]	(Lesquereux) Wolfe & Wehr	A sycamore	Macginitiea gracilis
Rosaceae	Amelanchier	Unidentified^[25]		A service berry Not described
Rosaceae	Neviusia	†Neviusia dunthornei^[26]	DeVore, Moore, Pigg, & Wehr	A snow wreath
Rosaceae	†Stonebergia	†Stonebergia columbiana^[27]	Wolfe & Wehr	A Sorbarieae genus
Sapindaceae	Acer	†Acer princetonense^[28]	Wolfe & Tanai	A maple
Sapindaceae	Acer	†Acer rousei^[28]	Wolfe & Tanai	A maple
Sapindaceae	Acer	†Acer stockeyae^[28]	Wolfe & Tanai	A maple
Sapindaceae	Acer	†Acer stewarti^[28]	Wolfe & Tanai	A maple
Sapindaceae	Acer	†Acer stonebergae^[28]	Wolfe & Tanai	A maple
Sapindaceae	Acer	†Acer toradense^[28]	Wolfe & Tanai	A maple
Sapindaceae	Acer	†Acer wehri^[28]	Wolfe & Tanai	A maple
Sapindaceae	Dipteronia	†Dipteronia brownii^[2]	McClain and Manchester	A Dipteronia species	Dipteronia brownii
Trochodendraceae	Tetracentron	†Tetracentron hopkinsii^[29]	Pigg et al.	A Tetracentron relative	Tetracentron hopkinsii
Trochodendraceae	Zizyphoides	Undescribed^[2]		A trochodendraceous species Not described
Ulmus	Ulmus	†Ulmus okanaganensis^[30]	Denk & Dillhoff	An elm	Ulmus okanaganensis
Urticaceae	Cf. Urticeae	Undescribed^[5]		A nettle not described to genus First identified as Rubus^[2]
Incertae sedis	†Chaneya	†Chaneya tenuis^[31]	(Lesquereux) Wang & Manchester	A sapindalean flower of uncertain affiliations	Chaneya tenuis

Mollusks

Family	Genus	Species	Authority	Notes
Hydrobiidae	†Micropyrgus	†Micropyrgus camselli^[32]	L.S. Russell, 1957	A hydrobiid mud snail
Lymnaeidae	Stagnicola	†Stagnicola tulameenensis^[32]	L.S. Russell, 1957	A lymnaeine pond snail
Physidae	Aplexa	†Aplexa ricei^[32]	L.S. Russell, 1957	An aplexine bladder snail
Physidae	Physa	†Physa saxarubrensis^[32]	L.S. Russell, 1957	A physine bladder snail
Planorbidae	Ferrissia	†Ferrissia arionoides^[32]	L.S. Russell, 1957	An ancylinine ramshorn snail
Planorbidae	Gyraulus?	Indeterminate^[32]	L.S. Russell, 1957	A possible planorbinine ramshorn snail Not described to species
Sphaeriidae	Sphaerium?	Indeterminate^[32]	L.S. Russell, 1957	A possible sphaeriine fingernail clam Not described to species

Insects

Coleopterans

Family	Genus	Species	Authority	Notes	Images
Cantharidae	Unidentified	Unidentified^[33]		A soldier beetle not identified to genus or species
Carabidae?	Unidentified	Unidentified^[33]		A caraboid superfamily beetle Displays traits similar to both Cicindelidae and Carabidae not identified to genus or species
Chrysomelidae	†Cryptocephalites	†Cryptocephalites punctatus^[34]	Scudder, 1895	A leaf beetle	Cryptocephalites punctatus (1895 illustration)
Chrysomelidae	Galerucella	†Galerucella picea^[35]	Scudder, 1879	A leaf beetle	Galerucella picea (1890 illustration)
Elateridae	Cryptohypnus?	†Cryptohypnus? terrestris^[35]	Scudder, 1879	A click beetle	Cryptohypnus? terrestris (1890 illustration)
Elateridae	Elaterites	Undescribed^[34]	Scudder, 1895	A click beetle Not described to species	Elateridae sp. indet (1890 illustration)
Elateridae	Limonius	†Limonius impunctus^[34]	Scudder, 1895	A wireworm click beetle	Limonius impunctus (1895 illustration)
Tenebrionidae	Tenebrio	†Tenebrio primigenius^[35]	Scudder, 1879	A darkling beetle	Tenebrio primigenius (1890 illustration)
Trogidae	Trox	†Trox oustaleti^[35]	Scudder, 1879	A hide beetle	Trox oustaleti (1890 illustration)

Dipterans

Family	Genus	Species	Authority	Notes	Images
Bibionidae	Penthetria	†Penthetria? fryi^[36]	Rice, 1959	A marchfly	Penthetria (?) fryi
Bibionidae	Penthetria	†Penthetria whipsawensis^[36]	Rice, 1959	A marchfly	Penthetria whipsawensis
Bibionidae	Plecia	†Plecia avus^[36]	(Handlirsch, 1910)	A marchfly First described as Penthetria avus (1910),^[37] moved to Plecia avus (1959)^[36]	Plecia avus
Bibionidae	Plecia	†Plecia canadensis^[36]	(Handlirsh, 1910)	A marchfly First described as Penthetria canadensis (1910),^[37] moved to Plecia canadensis (1959)^[36]	Plecia canadensis
Bibionidae	Plecia	†Plecia curtula^[36]	(Handlirsch, 1910)	A marchfly First described as Penthetria curtula (1910),^[37] moved to Plecia curtula (1959)^[36] Senior synonym of Penthetria avunculus (1959)	Plecia curtula
Bibionidae	Plecia	†Plecia dilatata^[36]	(Handlirsch, 1910)	A marchfly First described as Penthetria dilatata (1910),^[37] moved to Plecia dilatata (1959)^[36]	Plecia dilatata
Bibionidae	Plecia	†Plecia elatior^[36]	(Handlirsch, 1910)	A marchfly First described as Penthetria elatior (1910),^[37] moved to Plecia elatior (1959)^[36]	Plecia elatior
Bibionidae	Plecia	†Plecia minutula^[36]	Rice, 1959	A marchfly	Plecia minutula
Bibionidae	Plecia	†Plecia nana^[36]	(Handlirsh, 1910)	A marchfly First described as Penthetria nana (1910),^[37] moved to Plecia nana (1959)^[36]	Plecia nana
Bibionidae	Plecia	†Plecia pictipennis^[36]	(Handlirsh, 1910)	A marchfly First described as Penthetria pictipennis (1910),^[37] moved to Plecia pictipennis (1959)^[36] Penthetria lambei (1910), Penthetria ovalis (1910), & Penthetria separanda (1910) considered junior synonyms (1959)	Plecia pictipennis
Bibionidae	Plecia	†Plecia pulchra^[36]	(Handlirsch, 1910)	A marchfly First described as Penthetria pulchra (1910),^[37] moved to Plecia pulchra (1959)^[36]	Plecia pulchra
Bibionidae	Plecia	†Plecia pulla^[36]	(Handlirsh, 1910)	A marchfly First described as Penthetria pulla (1910),^[37] moved to Plecia pulla (1959)^[36] Penthetria brevipes (1910) considered a junior synonym (1959)	Plecia pulla
Bibionidae	Plecia	†Plecia reducta^[36]	(Handlirsh, 1910)	A marchfly First described as Penthetria reducta (1910),^[37] moved to Plecia reducta (1959)^[36]	Plecia reducta
Bibionidae	Plecia	†Plecia similkameena^[35]	(Scudder, 1879)	A marchfly First described as Penthetria similkameena (1879),^[35] moved to Plecia similkameena (1959)^[36]	Plecia similkameena
Bibionidae	Plecia	†Plecia transitoria^[36]	(Handlirsh, 1910)	A marchfly First described as Penthetria transitoria (1910),^[37] moved to Plecia transitoria (1959)^[36] Penthetria falcatula (1910) & Penthetria fragmentum (1910) considered junior synonyms (1959)	Plecia transitoria
Bibionidae	Plecia	†Plecia tulameenensis^[36]	Rice, 1959	A marchfly	Plecia tulameenensis
Dolichopodidae	†Microphor	†Microphor defunctus^[38]	(Handlirsch, 1910)	A long-legged fly First described as Microphorus defunctus (1910),^[37] moved to Microphor defunctus (1994)^[38]	Microphor defunctus (1910 illustration)
Ptychopteridae	†Etoptychoptera	†Etoptychoptera tertiaria^[39]	Handlirsh, 1909	A phantom cranefly	Etoptychoptera tertiaria (1910 illustration)
Tipulidae	Tipula	†Tipula tulameena^[37]	(Handlirsh, 1910)	A cranefly	Tipula tulameena

Hemipterans

Family	Genus	Species	Authority	Notes	Images
Aphrophoridae	Aphrophora	†Aphrophora angusta^[37]	Handlirsch, 1910	A spittlebug	Aphrophora angusta (1910 illustration)
Aphrophoridae	†Palaeoptysma	†Palaeoptysma venosa^[40]	Scudder, 1895	A spittlebug	Palaeoptysma venosa (1895 illustration)
Aphrophoridae	Palaphrodes	Undescribed^[40]	Scudder, 1895	A spittlebug Not described to species
Aphrophoridae	†Ptysmaphora	†Ptysmaphora fletcheri^[40]	Scudder, 1895	A spittlebug	Ptysmaphora fletcheri (1895 illustration)
Cercopidae	Cercopis	†Cercopis grandescens^[40]	Scudder, 1895	A froghopper	Cercopis grandescens (1895 illustration)
Cercopidae	Cercopis	†Cercopis selwyni^[35]	Scudder, 1879	A froghopper	Cercopis selwyni (1890 illustration)
Cercopidae	†Cercopites	†Cercopites torpescens^[40]	Scudder, 1895	A froghopper	Cercopites torpescens (1895 illustration)
Cercopidae	†Dawsonites	†Dawsonites veter^[40]	Scudder, 1895	A froghopper	Dawsonites veter (1895 illustration)
Cercopidae	Palecphora	Undescribed^[40]	Scudder, 1895	A froghopper Not described to species
Cercopidae	†Stenecphora	†Stenecphora punctulata^[40]	Scudder, 1895	A froghopper	Stenecphora punctulata (1895 illustration)
Cercopidae	†Stenolocris	†Stenolocris venosa^[40]	Scudder, 1895	A froghopper	Stenolocris venosa (1895 illustration)
Cicadellidae	Coelidia	†Coelidia columbiana^[35]	Scudder, 1879	A leafhopper	Coelidia columbiana (1890 illustration)
Fulgoridae	†Enchophora	Undescribed^[40]	Scudder, 1895	A fulgorid plant hopper Not described to species
Gerridae	†Telmatrechus	†Telmatrechus stali^[35]	(Scudder, 1879)	A gerrine water strider First described as Hygrotrechus stali (1879), moved to Telmatrechus stali (1895)^[40]	Telmatrechus stali (1890 illustration)
incertae sedis	†Planophlebia	†Planophlebia gigantea^[35]	Scudder, 1879	A hemipteran of uncertain placement	Planophlebia gigantea (1890 illustration)

Hymenopterans

Archibald, Mathewes, & Aase (2023) reported a Titanomyrma species ant queen from the Vermillion Bluffs site, and noted the range extension for Formiciinae into the highlands, as the subfamily was previously considered a strictly thermophilic ant group. Due to complications arising from preservational distortion during diagenesis, they were unable to determine the correct size of the queen in life. If the distortion was lateral, then compression to bilateral symmetry yielded an adult length of approximately 3.3 cm (1.3 in), placing it the same range as Formicium berryi and F. brodiei, known only from wings, and sugg4ested as possible males. Conversely stretching the fossil to bilateral symmetry results in a larger 5 cm (2.0 in) length estimate, placing it as comparable to queens of T. lubei and T. simillima.^[41]

Family	Genus	Species	Authority	Notes	Images
Braconidae	Bracon	Undescribed^[35]		A braconid wasp Not described to species	Bracon sp. (1890 illustration)
Formicidae	†Titanomyrma	Indeterminate^[41]		A formiciine titan ant Unplaced to species.	Titanomyrma sp.
Ichneumonidae	Xorides	†Xorides lambei^[37]	(Handlirsch, 1910)	A xoridine ichneumon parasitic wasp First named Xylonomus lambei (1910)^[37]	Xorides lambei
Tenthredinidae	Eriocampa	†Eriocampa tulameenensis^[42]	Rice, 1968	A tenthredinid sawfly	Eriocampa tulameenensis

Mecopterans

Family	Genus	Species	Authority	Notes	Images
†Dinopanorpidae	†Dinokanaga	†Dinokanaga wilsoni^[43]	Archibald, 2005	A dinopanorpid scorpion fly
Eomeropidae	†Eomerope	†Eomerope simpkinsae^[44]	Archibald & Rasnitsyn, 2018	An eomeropid scorpionfly

Neuropterans

Family	Genus	Species	Authority	Notes	Images
Ithonidae	†Ricaniella	†Ricaniella antiquata^[45]	(Scudder, 1895)	A Polystoechotid-group giant lacewing^[46] First described as Ricania antiquata (1895),^[40] moved to Ricaniella antiquata (1897)	Ricaniella antiquata (1895 illustration)

Odonata

Family	Genus	Species	Authority	Notes	Images
Aeshnidae	Indeterminate	Indeterminate^[47]		A daner dragonfly Wing too incomplete to determine genus affiliation.
Cf. †Dysagrionidae	†Allenbya	†Allenbya holmesae^[47]	Archibald & Cannings, 2022	A possible Dysagrionidae odonate. Not to be confused with the Princeton Chert waterlily Allenbya

Raphidiopterans

Family	Genus	Species	Authority	Notes	Images
Raphidiidae	†Megaraphidia	†Megaraphidia hopkinsi^[48]	Archibald & Makarkin, 2021	A Raphidiid snakefly

Vertebrates

Family	Genus	Species	Authors	Notes	Images
Amiidae	Cf. Amia	†"Amia" hesperia^[49]	Wilson, 1982	A bowfin	"Amia" hesperia scale
Aves incertae sedis	Unidentified	Unidentified^[50]	Mayr et al., 2019	Indeterminate feathers and a skeleton
Catostomidae	†Wilsonium	†Wilsonium brevipinne.^[51]^[52]^[53]	(Cope, 1893)	A catostomid sucker Originally described as Amyzon brevipinne Moved to Wilsonium in 2021
†Esthonychidae	†Trogosus	†Trogosus latidens^[54]^[55]	(Marsh, 1874)	A tillodont species
Hiodontidae	Hiodon	†Hiodon rosei^[51]^[56]	(Hussakof, 1916)	A mooneye First described as "Lucious" rosei (1916), moved to Eohiodon rosei in 1966, moved to Hiodon rosei in 2008^[56]
†Libotoniidae	†Libotonius	†Libotonius blakeburnensis^[51]	Wilson, 1977	A percopsiform fish
Salmonidae	†Eosalmo	†Cf. Eosalmo driftwoodensis^[51]	Wilson, 1977	An ancestral salmon
Trionychidae	Cf. Apalone	Undescribed^[57]		A soft shelled turtle Not described to species

Princeton Chert biota

The Princeton chert biota is unique in the Allenby formation due to the silicification of the chert, which has resulted in cellular and anatomical preservation of the organisms. As of 2016 over 30 different plant taxa had been described from chert fossils along with a number of fungal species.^[58]

Fungi

Order	Genus	Species	Authors	Notes
Ascomycota	†Cryptodidymosphaerites	†Cryptodidymosphaerites princetonensis^[59]	Currah, Stockey, LePage	An ascomycetan fungus on the host palm Uhlia allenbyensis
Ascomycota	†Monodictysporites	†Monodictysporites princetonensis^[60]	Klymiuk	An ascomycotan fungus hosted on Dennstaedtiopsis aerenchymata
Ascomycota	†Palaeoserenomyces	†Palaeoserenomyces allenbyensis^[59]	Currah, Stockey, LePage	An ascomycetan fungus on the host palm Uhlia allenbyensis

Ferns

Family	Genus	Species	Authors	Notes
Athyriaceae	†Dickwhitea	†Dickwhitea allenbyensis^[61]	Karafit et al.	An athyriaceous fern
Athyriaceae	†Makotopteris	†Makotopteris princetonensis^[62]	Stockey, Nishida, & Rothwell	An athyriaceous fern
Blechnaceae	†Trawetsia	†Trawetsia princetonensis^[63]	Smith et al.	A blechnacious fern
Dennstaedtiaceae	†Dennstaedtiopsis	†Dennstaedtiopsis aerenchymata^[64]	Cevallos-Ferriz, Stockey, & Pigg	A dennstaedtioid fern
Osmundaceae	Osmunda	Undescribed^[65]		An osmundaceous fern Not described

Conifers

Family	Genus	Species	Authors	Notes
Cupressaceae	Metasequoia	†Metasequoia milleri^[66]	Bassinger	A dawn redwood
Pinaceae	Pinus	†Pinus allisonii^[67]	Stockey	A 2-needled Pine foliage
Pinaceae	Pinus	†Pinus andersonii^[67]	Stockey	A 3-needled Pine foliage
Pinaceae	Pinus	†Pinus arnoldii^[68]^[69]	Miller	A basal Pine Cones belonging to the 5 needle Pinus similkameenensis foliage
Pinaceae	Pinus	†Pinus princetonensis^[67]	Stockey	A pinaceous cone
Pinaceae	Pinus	†Pinus similkameenensis^[68]^[69]	Miller	A basal Pine 5-needled foliage belonging to the Pinus arnoldii cones

Angiosperms

Family	Genus	Species	Authors	Notes
Alismataceae	†Heleophyton	†Heleophyton helobieoides^[70]	Erwin & Stockey	An aquatic or emergent water-plantain
Aponogetonaceae	Aponogeton	†Aponogeton longispinosum^[71]	Grímsson, Zetter, & Halbritter	A Cape-pondweed pollen
Araceae	†Keratosperma	†Keratosperma allenbyensis^[72]	Cevallos-Ferriz & Stockey	An arum family member
Arecaceae	†Uhlia	†Uhlia allenbyensis^[73]	Erwin & Stockey	A Coryphoid palm
Grossulariaceae	Ribes	Undescribed^[74]	Cevallos-Ferriz	A current fruit Not described
Lythraceae	Decodon	†Decodon allenbyensis^[75]	Cevallos-Ferriz & Stockey	A swamp loosestrife
Magnoliaceae	†Liriodendroxylon	†Liriodendroxylon princetonensis^[76]	Cevallos-Ferriz & Stockey	A Liriodendron-like wood.
Myrtaceae	†Paleomyrtinaea	†Paleomyrtinaea princetonensis^[77]	Pigg, Stockey & Maxwell	A Myrtaceous fruit
Nymphaeaceae	†Allenbya	†Allenbya collinsonae^[78]	Cevallos-Ferriz & Stockey	A water lily relative
Nyssaceae	Diplopanax	†Diplopanax eydei^[79]	Stockey, LePage, & Pigg	A tuplo relative.
Rosaceae	†Paleorosa	†Paleorosa similkameenensis^[80]	Bassinger	A rose family flower
Rosaceae	Prunus	†Prunus allenbyensis^[81]	Cevallos-Ferriz & Stockey	A prunoid wood.
Rosaceae	Prunus	"Species 1"^[81]	Cevallos-Ferriz & Stockey	A prunoid seed. Not described
Rosaceae	Prunus	"Species 2"^[81]	Cevallos-Ferriz & Stockey	A prunoid seed. Not described
Rosaceae	Prunus	"Species 3"^[81]	Cevallos-Ferriz & Stockey	A prunoid seed. Not described
Sapindaceae	†Wehrwolfea	†Wehrwolfea striata^[82]	Erwin & Stockey	A possible dodonaecous soapberry family flower
Saururaceae	Saururus	†Saururus tuckerae^[83]	Smith & Stockey	A lizard's-tail species
Vitaceae	Ampelocissus	†"Ampelocissus" similkameenensis^[84]^[85]	Cevallos-Ferriz & Stockey	A grape family fruit of uncertain generic placement^[85]
Vitaceae	incertae sedis	"Type 1"^[84]	Cevallos-Ferriz & Stockey	A grape family fruit of uncertain generic placement Not described
Vitaceae	incertae sedis	"Type 2"^[84]	Cevallos-Ferriz & Stockey	A grape family fruit of uncertain generic placement Not described
Cf. Iridaceae	†Pararisteapollis	†Pararisteapollis stockeyi^[86]	Zetter & Hesse	A possible iridaceous pollen morphotype
Incertae sedis	†Eorhiza	†Eorhiza arnoldii^[87]	Robison & Person	A semi-aquatic dicot of uncertain affinity.
Incertae sedis	†Ethela	†Ethela sargentiana^[88]	Erwin & Stockey	A cyperaceous or juncaceous monocot
Incertae sedis	†Princetonia	†Princetonia allenbyensis^[89]^[90]	Stockey	A possibly aquatic magnoliopsid flower of uncertain affiliation.
Incertae sedis	†Soleredera	†Soleredera rhizomorpha^[91]	Erwin & Stockey	A lilialean genus of uncertain placement

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[tarspot1998-59] Currah, R.S.; Stockey, R.A.; LePage, B.A. (1998). "An Eocene tar spot on a fossil palm and its fungal hyperparasite". Mycologia. 90 (4): 667–673. doi:10.1080/00275514.1998.12026955.

[Klymiuk2016-60] Klymiuk, A. A. (2016). "Paleomycology of the Princeton Chert. III. Dictyosporic microfungi, Monodictysporites princetonensis gen. et sp. nov., associated with decayed rhizomes of an Eocene semi-aquatic fern". Mycologia. 108 (5): 882–890. doi:10.3852/15-022. PMID 27302048. S2CID 7871220.

[dickwhitea2006-61] Karafit, S. J.; Rothwell, G. W.; Stockey, R. A.; Nishida, H. (2006). "Evidence for sympodial vascular architecture in a filicalean fern rhizome: Dickwhitea allenbyensis gen. et sp. nov.(Athyriaceae)". International Journal of Plant Sciences. 167 (3): 721–727. doi:10.1086/501036. S2CID 85348245.

[Makotopteris1999-62] Stockey, R. A.; Nishida, H.; Rothwell, G. W. (1999). "Permineralized ferns from the middle Eocene Princeton chert. I. Makotopteris princetonensis gen. et sp. nov.(Athyriaceae)". International Journal of Plant Sciences. 160 (5): 1047–1055. doi:10.1086/314191. PMID 10506480. S2CID 33465214.

[Trawetsia2006-63] Smith, S. Y.; Stockey, R. A.; Nishida, H.; Rothwell, G. W. (2006). "Trawetsia princetonensis gen. et sp. nov.(Blechnaceae): a permineralized fern from the Middle Eocene Princeton Chert". International Journal of Plant Sciences. 167 (3): 711–719. doi:10.1086/501034. S2CID 85160532.

[Review1991-64] Cevallos-Ferriz, S. R.; Stockey, R. A.; Pigg, K. B. (1991). "The Princeton chert: evidence for in situ aquatic plants". Review of Palaeobotany and Palynology. 70 (1–2): 173–185. Bibcode:1991RPaPa..70..173C. doi:10.1016/0034-6667(91)90085-H.

[Collinson2001-65] Collinson, M. E. (2001). "Cainozoic ferns and their distribution". Brittonia. 53 (2): 173–235. Bibcode:2001Britt..53..173C. doi:10.1007/BF02812700. S2CID 19984401.

[Metamilleri1981-66] Basinger, J. F. (1981). "The vegetative body of Metasequoia milleri from the Middle Eocene of southern British Columbia". Canadian Journal of Botany. 59 (12): 2379–2410. doi:10.1139/b81-291.

[Stockeypine1984-67] Stockey, R. A. (1984). "Middle Eocene Pinus remains from British Columbia". Botanical Gazette. 145 (2): 262–274. doi:10.1086/337455. S2CID 85063424.

[Miller1973-68] Miller Jr, C. N. (1973). "Silicified cones and vegetative remains of Pinus from Eocene of British Columbia". Contributions from the Museum of Paleontology, University of Michigan. 24: 101–118.

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[Heleophyton1991-70] Erwin, D. M.; Stockey, R. A. (1991). "Silicified monocotyledons from the Middle Eocene Princeton chert (Allenby Formation) of British Columbia, Canada". Review of Palaeobotany and Palynology. 70 ((1-2)): 147–162. Bibcode:1991RPaPa..70..147E. doi:10.1016/0034-6667(91)90083-F.

[Grímsson2014-71] Grímsson, F.; Zetter, R.; Halbritter, H.; Grimm, G. W. (2014). "Aponogeton pollen from the Cretaceous and Paleogene of North America and West Greenland: Implications for the origin and palaeobiogeography of the genus". Review of Palaeobotany and Palynology. 200 (100): 161–187. Bibcode:2014RPaPa.200..161G. doi:10.1016/j.revpalbo.2013.09.005. PMC 4047627. PMID 24926107.

[Keratosperma1988-72] Cevallos-Ferriz, S. R.; Stockey, R. A. (1988). "Permineralized fruits and seeds from the Princeton chert (Middle Eocene) of British Columbia: Araceae". American Journal of Botany. 75 (8): 1099–1113. doi:10.1002/j.1537-2197.1988.tb08822.x.

[Uhlia1995-73] Erwin, D.M.; Stockey, R.A. (1994). "Permineralized monocotyledons from the middle Eocene Princeton chert (Allenby Formation) of British Columbia: Arecaceae". Palaeontographica Abteilung B. 234: 19–40.

[Ribes1995-74] Cevallos-Ferriz, S. R. S. (1995). "Fruits of Ribes from the Princeton chert, British Columbia, Canada". American Journal of Botany. 82 (6).

[Decodon1988-75] Cevallos-Ferriz, S. R.; Stockey, R. A. (1988). "Permineralized fruits and seeds from the Princeton chert (Middle Eocene) of British Columbia: Lythraceae". Canadian Journal of Botany. 66 (2): 303–312. doi:10.1139/b88-050.

[Liriodendroxylon1990-76] Cevallos-Ferriz, S. R.; Stockey, R. A. (1990). "Vegetative remains of the Magnoliaceae from the Princeton chert (middle Eocene) of British Columbia". Canadian Journal of Botany. 68 (6): 1327–1339. doi:10.1139/b90-169.

[Paleomyrtinaea1993-77] Pigg, K. B.; Stockey, R. A.; Maxwell, S. L. (1993). ""Paleomyrtinaea", a new genus of permineralized myrtaceous fruits and seeds from the Eocene of British Columbia and Paleocene of North Dakota". Canadian Journal of Botany. 71 (1): 1–9. doi:10.1139/b93-001.

[Allenbya1989-78] Cevallos-Ferriz, S. R.; Stockey, R. A. (1989). "Permineralized fruits and seeds from the Princeton chert (Middle Eocene) of British Columbia: Nymphaeaceae". Botanical Gazette. 150 (2): 207–217. doi:10.1086/337765. S2CID 86651676.

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[Basinger1976-80] Basinger, JF (1976). "Paleorosa similkameenensis, gen. et sp. nov., permineralized flowers (Rosaceae) from the Eocene of British Columbia". Canadian Journal of Botany. 54 (20): 2293–2305. doi:10.1139/b76-246.

[Prunus1990-81] Cevallos-Ferriz, S. R.; Stockey, R. A. (1990). "Vegetative remains of the Rosaceae from the Princeton chert (Middle Eocene) of British Columbia". IAWA Journal. 11 (3): 261–280. doi:10.1163/22941932-90001183. S2CID 85023353.

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[Saururus2007-83] Smith, S. Y.; Stockey, R. A. (2007). "Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov.(Saururaceae) from the Middle Eocene Princeton Chert". American Journal of Botany. 94 (10): 1642–1657. doi:10.3732/ajb.94.10.1642. PMID 21636361.

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[Ethela1991-88] Erwin, D. M.; Stockey, R. A. (1992). "Vegetative body of a permineralized monocotyledon from the Middle Eocene Princeton chert of British Columbia". Courier Forschungsinstitut Senckenberg. 147: 309–327.

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Allenby Formation
Stratigraphic range: Ypresian 52.5–48 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Alternating Princeton Chert and coal sequences
Type	Geological formation
Unit of	Princeton Group, Eocene Okanagan Highlands
Sub-units	Princeton Chert, Vermillion Bluffs Shale
Overlies	Cedar Formation
Area	300 km² (120 sq mi)^[1]
Thickness	1,860–2,100 m (6,100–6,890 ft)^[1]
Lithology
Primary	Shale, sandstone
Other	Coal–breccia, coal–chert
Location
Coordinates	49°22.6′N 120°32.8′W / 49.3767°N 120.5467°W / 49.3767; -120.5467
Approximate paleocoordinates	53°06′N 107°30′W / 53.1°N 107.5°W / 53.1; -107.5
Region	British Columbia
Country	Canada
Extent	Princeton Basin & Tulameen basin
Type section
Named for	Allenby, British Columbia
Named by	Shaw
Year defined	1952
Allenby Formation (Canada) Show map of Canada Allenby Formation (British Columbia) Show map of British Columbia

Allenby Formation

Extent and correlation

Age

Lithology

Palynoflora

Compression paleobiota

Bryophytes

Pteridophytes

Gingkophytes

Pinophytes

Angiosperms

Mollusks

Insects

Coleopterans

Dipterans

Hemipterans

Hymenopterans

Mecopterans

Neuropterans

Odonata

Raphidiopterans

Vertebrates

Princeton Chert biota

Fungi

Ferns

Conifers

Angiosperms

References

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