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Princeton Chert

From Wikipedia, the free encyclopedia
Fossilized Flora
Close up of Princeton Chert outcrop showing volcanic ash (white layer at base), peaty coal (dark layer), and Chert layers (grey). Layer 36 is labelled.

ThePrinceton Chert is a fossil locality inBritish Columbia,Canada, which comprises ananatomically preserved flora ofEoceneEpoch age, with richspecies abundance and diversity. It is located in exposures of theAllenby Formation on the east bank of theSimilkameen River, 8.5 km (5.3 mi) south of the town ofPrinceton, British Columbia.[1]

History

[edit]

The Princeton Chert (Ashnola shale in older sources) and its fossils have been known since the 1950s,[2][3] but have attracted increased attention in the late 1970 and on.[4] This may be due to the rare type of silica permineralized fossilLagerstätten found, which has preserved plants and animals in minute 3D detail, with exceptional internal cellular detail.[5] This has meant anatomical descriptions and reconstruction of whole plants from isolated parts has been possible in many species.[6][7] Few plant fossils elsewhere in the world exhibit such excellence in both preservation and diversity. Similar aged fossil beds inEocene lake sediments are found elsewhere inBritish Columbia, including inDriftwood Canyon Provincial Park nearSmithers in northern British Columbia, theMcAbee Fossil Beds west ofKamloops, about 160 km (160,000 m) NNW of the Princeton Chert beds, and theKlondike Mountain Formation aroundRepublic, Washington, south of Princeton.

Location and geologic setting

[edit]

The Princeton Chert is an interbedded sequence consisting ofcoal,shale,volcanic ash, andchert in theAllenby Formation.[8][9][10][11] 49 chert layers, ranging in thickness from 1 to 55 cm (0.39 to 21.65 in) have been recognized and described, though each is not unique in organisms preserved. Despite this, trends are evident throughout theoutcrop, with certain taxa appearing and disappearing with time.[12][13]

The Princeton Chert was originally considered to beMiddle Eocene based on data frommammals, freshwater fish, and potassium-argon dates.[14][15][16] Recently, more accurateradiometric techniques provided a date of 48.7 mya,[12] placing the Princeton Chert in theYpresian stage (47.8–56.0 mya), consistent with the wholeAllenby Formation being now dated radiometrically as beingEarly Eocene.[10][17][18]

Theclimate at this time was warm; it had reached a maximum during a series of warming events during the Early Eocene with the Princeton Chert likely deposited after theEocene Thermal Maximum 2 and during the Early Eocene Climatic Optimum.[10][19][20][21] During this time the sea warmed approximately 4 °C and terrestrial temperatures were several degrees warmer than today, meaning little or no ice was present at the poles. The temperature difference between poles and equator was small.[22][23] This long term warmth is thought to be due to increased greenhouse gases, particularly CO2 trapping more heat.[21][24] The reason for this sudden increase in CO2 is unknown, but it is hypothesised that it was due to an increase in ocean floor being recycled via volcanic arcs andmetamorphic decarbonation reactions.[23] This happened because the ocean between India and Asia was disappearing and being replaced by theHimalayas and theTibetan Plateau due to the collision of tectonic plates. Also at the time,Australia, which was joined toAntarctica, was beginning to move northwards.[25]

The Princeton Chert fossils indicate that the area was anaquatic ecosystem, growing intropical tosubtropical conditions.[12] More recent analysis of the fossil flora, however reconstructs for the Princeton Chert flora a moist warm temperate climate with mean annual temperature 13.1 ± 3.1 °C, with mild winters (cold month mean temperature 5.3 ± 2.8 °C), and mean annual precipitation 114 ± 42 cm per year.[20] Several of the smaller chert layers are separated byvolcanic ash layers, indicating nearby volcanic activity. It is thought that fossils were pervaded with silicic acid due to this volcanic activity.[26] Subsequently, water charged with minerals flowed from springs orgeysers into the low lying basin where the Princeton chert was located. Here, the water surrounded organisms as they grew, along with plant debris which had been accumulated.[12] Many organisms were preserved in situ, in the lake or small pond environment in which they lived.[27] The preservation must have been rapid, due to the minute cellular detail which has been conserved. This sequence of events is thought to have been replicated up to 50 times, as the basin allowed peat to re-accumulate each time,[12] producing the multiple layers.

Known biota

[edit]

Sampling into the Princeton Chert has been carried out, but presently the data has not been analysed in detail.[12] Across the outcrop, trends in taxa can be seen; in the topmost layers fossil organs ofMetasequoia milleri[9] cease to be represented, yetPinus (pine) and monocotyledons increase in number. There is a huge increase inferns, such asDennstaedtiopsis, after a huge ash fall, though few angiosperms occur in these layers. A large number of angiosperms have been found along with several types ofconifers, ferns, and several unidentified fossils from various families.[28]

In situ lacustrine fossils

[edit]

The array of floral and faunal fossils found in the Princeton Chert has offered unequivocal evidence that it was alacustrine or lake environment. The plant fossils found show many structural and anatomical adaptations to an aquatic environment, including a reduced vascular system,aerenchyma in tissues (air spaces to provide buoyancy), andprotoxylem lacunae surrounded by a ring of cells with thickened inner walls.[27][12] Further evidence is provided by the fossils’ clear affinities with modern aquatic angiosperms. Many extant plants show these adaptations and are similar to the organisms found in the chert. For example, water lilies (Allenbya,Nymphaeaceae), water plantains (Alismataceae),[29] arums (Keratosperma,Araceae)[6][30] and rushes and sedges (Ethela,Juncaceae/Cyperaceae) are just some of the angiosperms found both today and in the Princeton Chert. Seeds have also been found which share adaptations with living aquatics.[27][12] On the other hand, terrestrial fossils have rarely been found. The few that are, are represented mainly by seeds, some of which may have been transported by birds.[9][28][31]

Additional support for the aquatic nature of the Princeton Chert deposits comes from animal fossils. Several fossils of a freshwater fish,Amia (bowfin), have been found in the shale overlying the plant deposits, along with remains of the freshwater fishesAmyzon andLibotonius, plus a soft-shelled turtle.[27][32]

Fungi

[edit]

Pathogenicfungi have been recorded on the leaves and other organs of some vascular plants. FossilUhliapalms have tar spot fungi on their leaves described asPaleoserenomyces, which is in turn parasitized by amycoparasite,Cryptodidymosphaerites princetonensis.[33] Symbioticmycorrhizal relationships have also been preserved in roots ofPinus andMetasequoia milleri. InMetasequoia these associations have been compared to extantmycorrhizae, and found to be very similar.[12] The mycorrhizal relationship withPinus was the first documentation ofectomycorrhizae from the fossil record, with the fungi suggested as close to the modern pine symbioticsRhizopogon andSuillus[34]

Paleofauna

[edit]
FamilyGenusspeciesNotesImages

Undescribed

Undescribed

Undescribed[35]

A lepidopteran forewing
Not described to family/genus/species

Undescribed

Undescribed

undescribed[36]

Aturtle,
Bones preserved in the Princeton Chert

Paleoflora

[edit]

Pteridophytes

[edit]
FamilyGenusspeciesNotesImages
Athyriaceae

†Dickwhitea

†Dickwhitea allenbyensis[37]

Anathyriaceous fern

†Makotopteris

†Makotopteris princetonensis[38]

Anathyriaceous fern

Blechnaceae

†Trawetsia

†Trawetsia princetonensis[39]

Ablechnacious fern

†Dennstaedtiopsis

†Dennstaedtiopsis aerenchymata[40]

Adennstaedtioid fern

Osmundaceae

Osmunda

Undescribed[41][20]

Anosmundaceous fern
Not identified to species

Conifers

[edit]
FamilyGenusSpeciesNotesImages

Cupressaceae

Metasequoia

†Metasequoia milleri[42]

Adawn redwood

PinaceaePinus

†Pinus allisonii[43]

A 2-needledPine

†Pinus andersonii[43]

A 3-needledPine

†Pinus arnoldii[44][45]

A basalPine,
First described for ovulate cones
A whole plant reconstruction
includes the synonymizedP. similkameenensis.[45]

†Pinus princetonensis[43]

Apinaceous cone

Angiosperms

[edit]
FamilyGenusSpeciesNotesImages

Alismataceae

†Heleophyton

†Heleophyton helobieoides[46]

An aquatic or emergentwater-plantain

Aponogetonaceae

Aponogeton

†Aponogeton longispinosum[47]

ACape-pondweed pollen

Araceae

†Keratosperma

†Keratosperma allenbyensis[6]

Alasioidarum family seed genus[30]

Arecaceae

†Uhlia

Uhlia allenbyensis[48]

ACoryphoid palm

Grossulariaceae

Ribes

Undescribed[49]

Acurrent fruit
Not described

Cf.Iridaceae

†Pararisteapollis

†Pararisteapollis stockeyi[50]

A possibleiridaceous pollen morphotype

Lauraceae

Undescribed

Undescribed[51]

Alauraceous fruit.
Briefly described by Littleet al (2009) but not named

Lythraceae

Decodon

†Decodon allenbyensis[52]

Aswamp loosestrife
Initially described from seeds
Little & Stockey (2003) provided a whole plant reconstruction[53]

Magnoliaceae

†Liriodendroxylon

†Liriodendroxylon princetonensis[54]

ALiriodendron-like wood.

Myrtaceae

†Paleomyrtinaea

†Paleomyrtinaea princetonensis[55]

AMyrtaceous fruit

Nymphaeaceae

†Allenbya

†Allenbya collinsonae[56]

Awaterlily relative.
Not to be confused with the odonateAllenbya[57]

Nyssaceae

Diplopanax

†Diplopanax eydei[58]

Atuplo relative.

Rosaceae

†Paleorosa

†Paleorosa similkameenensis[8]

A rose family flower

Prunus

†Prunus allenbyensis[59]

Aprunoid wood.

"Princeton chert species 1"[59]

Aprunoid seed.
Not described to species

"Princeton chert species 2"[59]

Aprunoid seed.
Not described to species

"Princeton chert species 3"[59]

Aprunoid seed.
Not described to species

Sapindaceae

†Wehrwolfea

†Wehrwolfea striata[60]

A possibledodonaecoussoapberry family flower

Saururaceae

Saururus

†Saururus tuckerae[61]

Alizard's-tail species

Vitaceae

Ampelocissus

†"Ampelocissus" similkameenensis[62][63]

Agrape family fruit of uncertain generic placement[63]

incertae sedis

"Type 1"[62]

Agrape family fruit of uncertain generic placement
Not described

"Type 2"[62]

Agrape family fruit of uncertain generic placement
Not described

incertae sedis

†Eorhiza

†Eorhiza arnoldii[64][36]

A semi-aquaticdicot of uncertain affinity.

†Ethela

†Ethela sargentiana[65]

Acyperaceous orjuncaceouspoalean monocot

†Princetonia

†Princetonia allenbyensis[13][66]

A possibly aquaticmagnoliopsid
flower of uncertain affiliation.

†Soleredera

†Soleredera rhizomorpha[67]

Alilialean genus of uncertain placement

Fungi

[edit]
OrderGenusspeciesNotesImages

Dothideales

†Palaeoserenomyces

†Palaeoserenomyces allenbyensis[68]

Anascomycetan fungus on the host palmUhlia allenbyensis

Jahnulales

Xylomyces

undescribed[69]

Ajahnulalean fungi.
Noted to be similar toXylomyces giganteus.
In situ decomposer ofEorhiza arnoldii

Microascales

Culcitalna

undescribed[69]

Amicroascalean fungi.
Noted to be similar toCulcitalna achraspora.
In situ decomposer ofEorhiza arnoldii

Thielaviopsis

undescribed[69]

Amicroascalean fungi.
Noted to be similar toThielaviopsis basicola.
In situ decomposer ofEorhiza arnoldii

Pleosporales

†Cryptodidymosphaerites

†Cryptodidymosphaerites princetonensis[68]

Anascomycetan fungus
hosted onUhlia allenbyensis

†Monodictysporites

†Monodictysporites princetonensis[70]

Anascomycotan fungus
hosted onDennstaedtiopsis aerenchymata

Undescribed

Undescribed

Ectomycorrhizae fungi associated withPinus roots
similar toRhizopogon andSuillus
Not described to genus or species[34]

See also

[edit]

References

[edit]
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  2. ^Arnold, C. A. (1955). "A Tertiary Azolla from British Columbia".Contributions from the Museum of Paleontology, University of Michigan.12 (4):37–45.
  3. ^Boneham, R.F. (1968). "Palynology of three Tertiary coal basins in south-central British Columbia".Ph.D. Dissertation, University of Michigan, Ann Arbor, Michigan.
  4. ^Greenwood, D.R.; Pigg, K.B.; Basinger, J.F.; DeVore, M.L. (2016)."A review of paleobotanical studies of the Early Eocene Okanagan (Okanogan) Highlands floras of British Columbia, Canada and Washington, USA".Canadian Journal of Earth Sciences.53 (6):548–564.Bibcode:2016CaJES..53..548G.doi:10.1139/cjes-2015-0177.hdl:1807/71961.
  5. ^Pigg, KB; Stockey, RA (1996). "The significance of the Princeton chert permineralized flora to the middle Eocene upland biota of the Okanogan Highlands".Washington Geology.24:32–36.
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  70. ^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.

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