User:Kevmin/sandbox/Palaeocarpinus
| Kevmin/sandbox/Palaeocarpinus Temporal range: Early Paleocene?-Early Oligocene?
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| Palaeocarpinus sp. fruit | |
Scientific classification 
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| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Rosids |
| Order: | Fagales |
| Family: | Betulaceae |
| Subfamily: | Coryloideae Manchester |
| Genus: | †Palaeocarpinus P.R. Crane |
| Type species | |
| Palaeocarpinus laciniata P.R. Crane
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| Species | |
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Palaeocarpinus is an extinct genus of plants placed in the Betulaceae subfamily Coryloideae. The genus had a wide spread circumboreal distribution, with species found in North America, Europe, and Asia. The first appearance dates to the Early Paleocene and species are spread though the Paleocene and Eocene with the youngest species occurring at the Eocene-Oligocene boundary. A total of thirteen species have been named with a small grouping of referred specimens that have not been redescribed in the modern literature. The first species to be described were placed into several unrelated genera or into a form taxon for fruits. Palaeocarpinus is now placed basally into Coryloideae, with an unresolved relationship to the two modern coryloid tribes Coryleae and Carpinieae.
Distribution and age
[edit]Palaeocarpinus species have a Circumboreal distribution across North America, Europe, and Asia. The ages of the species are concentrated in the early Paleogene, with most species being Paleocene to Eocene aged, though the youngest is possibly placed just into the Oligocene.[1] Two occurrences of the genus in Europe date from the PAleocene, while the third is tentatively identified as Eocene. P. laciniata are type localitied in strata of the Reading Formation outcropping in the Cold Ash quarry, near Newbury, Berkshire England. While the exact age is uncertain, the general age is late Paleocene (upper Paleocene of Crane, 1981).[2] To the south in France, fossils recovered from the Menat Formation near Menat in Puy-de-Dôme have an agreed upon age of late Paleocene based on associated fossils and volcanic deposits.[1] The northernmost occurrence of Palaeocarpinus are from the arctic Spitzbergen in western Svalbard. The fossils are produced by strata of the Storvola Formation which Golovneva (2002) reported outcropping along Adventfjorden,[3] the site was later reported being on Prins Karls Forland by Correa-Narvaez and Manchester however. The exact age is uncertain, but based on surrounding stratigraphy plus palynologiacl data, its most often cited to be Eocene.[1]
Of the three Asian occurrences divided between Russia with two species and China with one, and includes both the oldest and youngest occurrences for the genus[1] The western most site is south of the Irtysh river in the Altay Prefecture from rocks of the Wulungu Depression. Based on the associated plant flora of the formation, a Paleocene age is applied to the formation.[4] The older of the two Russian localities, from which P. pacifica originates, is located near the lower course of the Amur in Khabarovsk Krai in the Russian Far East. The Malo-Mikhailovka Formation which outcrops near Puer Ridge between the Amur and the Strait of Tartary, has been identified as from the Early Paleocene's Danian age.[5] In contrast, the Sikhote-Alin site on the Strait of Tartary's Buoy Bay, 8 km (8,000 m) south of Nelma village is the youngest of the known Palaeocarpinus sites. The locality is part of the Kizi Group, with the site included in the uppermost strata of the Buoy Bay Member. Floral and mammalian correllaitions are indestinct and give possible ages between the Late Eocene and the Early Oligocene. One volcanic rock yielded a Potassium-argon date of 34 ± 0.8 million years old, within a million years of the Eocene-Oligocene boundary.[6]
The greatest concentration of species and localities is found across the Western regions of North America, with species groups in the Great Plains-Rocky Mountains, in the Oregon interior, and in the greater Okanagan region of British Columbia and Washington. The northern most of the great plains localities is at Munce's Hill near Joffre, Alberta where Paskapoo Formation rocks of possible Lacombe member affinity outcrop in a road cut. The age of the Lacombe member is accepted to likely be Late Paleocene in the Tiffanian.[7] Most of the Great Plains and Rocky Mountains sites are plced into various Fort Union Group formations. The two western North Dakota occurrences, at Almont and Beicegel Creek respectively, are both in the Tiffanian Sentinel Butte Formation of the Fort Union Group,[8] while the eastern Montana sites around Big Timber are Fort Union Formation.[1] All species in the Wyoming region are known from Fort Union Formation fossils, with most sites in Wyoming in the P-3 pollen zone and thus Middle Paleocene.[9][1]
Palaeocarpinus specimens have been recovered from at least twelve individual sites in the Okanagan highlands, with named species from the Coldwater Beds near Quilchena[10] and Allenby Formation near Princeton, British Columbia, plus the Klondike Mountain Formation in Republic, Washington, northern Ferry County, Washington.[11][1] The Okanagan highlands are aged between 51.5 ± 0.4 million years ago for the Quilchena site to 51.2 to 49.4 million years ago for the Klondike Mountain Formations Tom Thumb Tuff member.[12] Additional Palaeocarpinus sp. fossils were reported from Okanagan highlands sites of the Tranquille Formation near Cache Creek, British Columbia and the Horsefly Shales near Horsefly, British Columbia in 1908.[10] Pigg, Manchester, and Wehr added the report of unplaced fruits in the Driftiwood Shales near Smithers, British Columbia, the Golden Promise site outside Republic, and Corkscrew Mountain near Toroda, Washington.[11]
All four of the fossil sites in Oregon are placed within the Clarno Formation outcropping in the Blue Mountains area in the northeastern part of the state. Several methods of radiometric dating have been applied to that unit of the Clarno, with coinciding results. Fission track data from zircon crystals and 40Ar/39Ar data from plagioclase crystals have both yielded an Eocene age between 46.8 to 44.7 MYA, placing the fossils as from the Lutetian.[1]
History & classification
[edit]Around the same time as the Work with Menat fossils was happening, the first geologic explorations of was will become British Columbia were documenting the Okanagan Highlands. Carpolithes dentata specimens collected during the 1888 expedition to the Stump lake area were listed as included in the Redpath Museum collections by George Mercer Dawson (1890).[13]
Description
[edit]All Palaeocarpinus species fruits have a small Carpinus like nutlets with ribbing running length-wise on the surface. They all also have a pair of surrounding bracts with varying degrees of specialization in shape and size. The fruits are born in cymule pairs on long infructescences to which some species fruits detach at maturity while others remain affixed as a group.[1]
Palaeocarpinus aspinosa Manchester & Chen
[edit]Palaeocarpinus barksdaleae Pigg, Manchester & Wehr
[edit]
One Palaeocarpinus barksdaleae fruiting head has been described, with a mass of approximately ten fruits grouped helically along the central stem. Fruits are most frequently found alone, rather than in a pair or as a group on stem, likely do to dispersal as lone fruits being a part of the reproductive strategy. The nutlets are enclosed between two unlobed, fan shaped bracts. The bract tissue is thickened and there are around twelve palmate veins which branch towards the upper margins. The general size of the bracts ranges between 4.5–10 mm (0.18–0.39 in) tall and 6–11 mm (0.24–0.43 in) wide giving an almost square outline. Some of the secondary veins, and all of the primaries, terminate in the bract spines, while a network of fine reticulate tertiary veins interlaces between the larger veins as well as looping back on themselves in the bract margin. The small nutlets have a flattened base and a slightly elongated rounded outline with eight to ten total ribs arising around the nutlet. The nutles are only 1.3–3.2 mm (0.051–0.126 in) wide and between 2.5–5 mm (0.098–0.197 in) tall and the nut tip has a 1 mm (0.039 in) tall style that possibly splits in two near the tip.[11][1]
The species was described by Kathleen Pigg, Manchester & Wesley Wehr (2003) from a holotype and a series of 89 paratypes. The holotype, UWBM 97408A & B, and 82 of the paratypes were included in the Burke Museum of Natural History and Culture (UWBM) palaeobotanical collections. A small group of five paratypes were placed in the Arizona State University collections, while the final two were University of Florida specimens. The type locality within the Klondike Mountain Formation was identified as UWBM site B5077 - "Gold Mountain". Other sites of the Klondike Mountain Formation have also produced P. barksdaleae fruits, including UWBM B4131 "Boot Hill", B2737 "Knob Hill" and UWBM B1795 "Resner Canyon". The specific epithet barksdaleae was designated as a matronym honoring Lisa Barksdale in recognition of the 14 years she spent as curator for the Stonerose Interpretive Center, guiding public and scientific access to the Republic fossils.[11]
Palaeocarpinus borealis (Heer) Correa & Manchester
[edit]The earliest reported fossils now attributed to Palaeocarpinus were from the Menat Formation, with Oswald Heer (1859) publishing a group of fossils as Anchietea borealis. Nearly 20 years later, another fossil from Menat was given the name Corylus lamottii by Gaston de Saporta (1877), but no description was supplied with the name so it was considered nomen nudum. Heers' Menat fossils were later redescribed and moved to Atriplex borealis by Louis Laurent. While this placement was unchanged until the 1981 publication of the genus Palaeocarpinus by Peter Crane, the affinity of the fossils to Betulaceae was first suggested by Saporta and Antoine-Fortuné Marion (1885), who put forward the connection between the fruits and leaves called Corylus mac-quarii. Crane noted in his work on the English fossils that the Menat specimens were also from the same genus, but did not present a formal reassignment. His change was accepted by later workers, but an official redescription was not presented until Julian E. Correa-Narvaez and Steven Manchester (2021) in their monograph on the genus. Over the preceding century Heers original specimen, which he did not illustrate or note as holotype, was lost. As such, Correa-Narvaez & Manchester chose to designate "MEN 19" in the National Museum of Natural History, France as neotype.[1]
Palaeocarpinus dakotensis Manchester, Pigg & Crane
[edit]Palaeocarpinus dentata (Penhallow) Pigg, Manchester & Wehr
[edit]The lectotype fruit has broad shallowly toothed bracts that are 10 mm (0.39 in) wide and 8 mm (0.31 in) tall.[11][1]
Palaeocarpinus dentata is the first Okanagan highlands species described, being first discussed by David P. Penhallow in 1890. The original fruit was collected in 1888 by George Mercer Dawson during summer field work in the region. Found near Stump Lake in the Quilchena, British Columbia area Dawson very briefly detailed the specimen and noted that it was superficially similar to Carpinus nutlets, but not referable to the genus. The bracts of the specimen are notably faint on the fruit and Dawson attributed it to being displaced pericarp, outer fleshy fruit tissues. Penhallow chose to name the species Carpolitus dentatus, the genus placement being a form genus for plant fruits and seeds of uncertain affinity. The specimen was deposited in the Redpath Museum by the Geological Survey of Canada as specimen 2.2420.[13] The fossils from sStump Lake are part of the Coldwater Beds, and the fossils are realted to the near by Quilchena flora. The specimen was later designated the lectotype for the species by Pigg, Manchester, and Wehr (2003), and they moved the species from Carpolithus to Palaeocarpinus dentatus. They noted that the specific epithet dentata was likely a reference to the toothed outline of the bracts.[11] The species name was further corrected by Correa and Manchester who noted Palaeocarpinus is a feminine Latin noun in construction, so the species name should have been shifted to P. dentata when moved from Carpolithus which is a neuter noun.[1]
Palaeocarpinus joffrensis Sun & Stockey
[edit]Palaeocarpinus laciniata Crane
[edit]Palaeocarpinus orientalis Manchester & Guo
[edit]Palaeocarpinus pacifica Akhmetiev & Goloveneva
[edit]Palaeocarpinus parva Correa-Narvaez & Manchester
[edit]Palaeocarpinus pterabaratra Correa & Manchester
[edit]Palaeocarpinus pteravestigia Correa & Manchester
[edit]Palaeocarpinus sikhotealinensis Akhmetiev & Manchester
[edit]Palaeocarpinus stonebergae Pigg, Manchester & Wehr
[edit]P. stonebergae was initially described from only singular fruits, but a few incomplete infructescences have since been discovered. The known sections are between 3.3–7.5 mm (0.13–0.30 in) wide and 13.8–14.6 mm (0.54–0.57 in) long and bear the densely packed fruits as pairs between short internodes, with a total of 11 fruits on the largest infructescence fossil. The 3–8 mm (0.12–0.31 in) tall by 3.4–14 mm (0.13–0.55 in) wide individual fruits are accompanied by a pair of deeply lobed spiny bracts with highly reduced membrane. The major veins run up the approximately ten narrow lobes, which occasionally fork, and frequently end in at the spiny tips. As with other species, the rounded elliptical nutlet has a forked style at the nut tip and has four to eight ribs. The nutles are between 1.5–4.5 mm (0.059–0.177 in) wide and 1.1–5 mm (0.043–0.197 in) tall. Based on the notably reduced membrane, spiny lobe tips and preservation as single fruits the means of dispersal was possibly clinging to passing animals.[11][1]
Palaeocarpinus stonebergae is another Okanagan highlands species described by Pigg, Manchester & Wehr (2003), this one from the Princeton, British Columbia area. The type series includes the holotype, UWBM 77466, and a series of 22 paratypes in the Burke Museum, plus one paratype at the University of Florida. The type locality within the Allenby Formation was UWBM site B4294 - "Thomas Ranch" while only one other Princeton area site, UWBM site B3264 "Coalmont Bluffs" was noted to have produced specimens. The specific epithet stonebergae in a matronym honoring Margaret Stoneberg for her work bringing scientific and public attention back to the fossils of the Princeton region.[11]
Undescribed Palaeocarpinus fossils
[edit]In the material collected by Lawrence Lambe's 1908 collecting expedition though central British Columbia, David P. Penhallow reported additional Carpolithes dentata fossils. Penhallow deemed the three Horsefly, two Quilchena, and two or three Tranquille River fossils as all equivalent to Dawsons original Carpolithes dentata specimen. In total between seven and eight additional fossils were noted, but have been largely ignored since the original mention by Penhallow.[10] Pigg, Manchester, and Wehr added the report of unplaced fruits in the Driftiwood Shales near Smithers, British Columbia, the Golden Promise site, UWBM B4876, Outside Republic, and Corkscrew Mountain, UWBM B6494, near Toroda, Washington. They considered this fruits as not placeable in the species known in the area at the time, and did not name new species from them.[10] None of the unplaced fossils noted in 2003 were discussed by Correa and Manchester (2021).
Paleoenvironment
[edit]The distribution of Palaeocarpinus species follows a similar trend to that seen in the living species of Corylus and Carpinus. Fossil sites are usually from an upland temperate or humid environment along lakes, streams, or rivers.[1] The Rocky Mountain and Great Plains sites are all from various river system environments such as flood plain, point bar, and overbank areas. These rivers were likely affiliated with the ongoing Laramide Orogeny resulting in mountain building across Western North America. The Okanagan Highlands preserves a series of upland lake system surrounded by a mixed conifer–broadleaf forests with nearby volcanism.[14] Sites in the Okanagan Highlands are grouped around a series of upland temperate lakes stretching 1,000 km (1,000,000 m) southwest from Central British Columbia to Northeast central Washington. Volcanics from the Challis Arc and associated uplift and graben formation characterize the lakes. The sites in Oregon are a mix of volcanics and riverine systems, with the fossils associated with river or lake areas where fish were present.[15][1]
The European sites are from a mix of depositional environments. The Cold Ash Quarry site shows typical silt lenses of clays and silts plus cross bedding that are all indicators of a river system. In contrast, the Menat fossils are preserved in a maar, a shallow lake environment in an infilled volcanic crater. The waters of the lake were mineral rich and preserved the flora from the surrounding crater rim hills. To the north, the Spitzbergen site has a mix of shales and sandstones with associated plant and mollusk fossils suggesting an estuary or lake delta environment.[1]
The three Asian sites are not as well defined as the others, with only the Puer Ridge site generally explored. While both the Alti and Buoy Bay sites are in silt and claystones, the depositional nature of them is unreported. With the Bouy Bay site the locality is noted to have likely been a higher elevation or warm temperate site, and the Alti was likely a humid temperate environment with a deciduous forest close to a coast. Puer Ridge was reported to be an intermountain depression with areas of swamp.[1]
References
[edit]- ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac Correa-Narvaez, J. E.; Manchester, S. R. (2021). "Distribution and Morphological Diversity of Palaeocarpinus (Betulaceae) from the Paleogene of the Northern Hemisphere". The Botanical Review. 88 (2): 161–203. doi:10.1007/s12229-021-09258-y. S2CID 237795532.
- ^ a b Crane, P.R. (1981). "Betulaceous leaves and fruits from the British Upper Palaeocene". Botanical Journal of the Linnean Society. 83 (2): 103–136. doi:10.1111/j.1095-8339.1981.tb01224.x.
- ^ a b Golovneva, L.B. (2002). "Palaeocarpinus (Betulaceae) from the Paleogene of Spitsbergen and transatlantic floristic migrations". Paleontological Journal. 36 (4): 422–428.
- ^ a b Manchester, S.R.; Guo, S.-X. (1996). "Palaeocarpinus (extinct Betulaceae) from northwestern China: new evidence for Paleocene floristic continuity between Asia, North America, and Europe". International Journal of Plant Sciences. 157 (2): 240–246. doi:10.1086/297343.
- ^ a b Akhmet'ev, M.A.; Golovneva, L.B. (1998). "New data on composition and age of Malomikhailovka flora from the Upper Cretaceous of the Amur River lower courses". Stratigraphy and Geological Correlation. 6 (3): 249–261.
- ^ a b Akhmetiev, M.A.; Manchester, S.R. (2000). "A new species of Palaeocarpinus (Betulaceae) from the Paleogene of Eastern Sikhote-Alin". Paleontological Journal. 34: 467–474.
- ^ a b Sun, F.; Stockey, R.A. (1992). "A new species of Palaeocarpinus (Betulaceae) based on infructescences, fruits, and associated staminate inflorescences and leaves from the Paleocene of Alberta, Canada". International Journal of Plant Sciences. 153 (1): 136–146. doi:10.1086/297015.
- ^ a b Manchester, S.R.; Pigg, K.B.; Crane, P.R. (2004). "Palaeocarpinus dakotensis sp. n. (Betulaceae: Coryloideae) and associated staminate catkins, pollen, and leaves from the Paleocene of North Dakota". International Journal of Plant Sciences. 165 (6): 1135–1148. doi:10.1086/423870.
- ^ a b Manchester, S.R.; Chen, Z.D. (1996). "Palaeocarpinus aspinosa sp. nov. (Betulaceae) from the Paleocene of Wyoming, USA". International Journal of Plant Sciences. 157 (5): 644–655. doi:10.1086/297386.
- ^ a b c d Penhallow, D. P. (1908). Report on Tertiary plants of British Columbia collected by Lawrence M. Lambe in 1906 together with a discussion of previously recorded tertiary floras (Report). Geological series; Contributions to Canadian Paleontology. Ottawa, Canada: Geological Survey of Canada. pp. 1–167.
- ^ a b c d e f g h Pigg, K.B.; Manchester S.R.; Wehr W.C. (2003). "Corylus, Carpinus, and Palaeocarpinus (Betulaceae) from the Middle Eocene Klondike Mountain and Allenby Formations of Northwestern North America". International Journal of Plant Sciences. 164 (5): 807–822. doi:10.1086/376816. S2CID 19802370.
- ^ Rubino, E.; Leier, A.; Cassel, E.; Archibald, S.; Foster-Baril, Z.; Barbeau, D. Jr (2021). "Detrital zircon UPb ages and Hf-isotopes from Eocene intermontane basin deposits of the southern Canadian Cordillera". Sedimentary Geology. 105969 (online).
- ^ a b Dawson, J. W. (1890). On fossil plants from the Similkameen Valley and other places in the southern interior of British Columbia. Royal Society of Canada.
- ^ Archibald, S.; Greenwood, D.; Smith, R.; Mathewes, R.; Basinger, J. (2011). "Great Canadian Lagerstätten 1. Early Eocene Lagerstätten of the Okanagan Highlands (British Columbia and Washington State)". Geoscience Canada. 38 (4): 155–164.
- ^ "Clarno Assemblage". national Park Service - John Day Fossil Beds National Monument Oregon. Retrieved August 29, 2024.
External links
[edit]
Media related to Palaeocarpinus at Wikimedia Commons
† Category:Prehistoric angiosperm genera Category:Fossil taxa described in 1981 Category:Eocene life of North America Category:Eocene plants Category:Flora of Oregon Category:Extinct flora of North America Category:Prehistoric plants of North America Category:Allenby Formation Category:Clarno Formation Category:Klondike Mountain Formation Category:Fossils of British Columbia