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Kevmin/sandbox/Bohlenia
Temporal range: 51–49 Ma
↓
 Early Eocene (Ypresian)
Bohlenia americana lectotypes
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Sapindales
Family: Sapindaceae
Subfamily: incertae sedis
Genus: †Bohlenia
Wolfe & Wehr
Species
  • Bohlenia americana
  • Bohlenia insignis
Synonyms
B. americana synonymy
  • Comptonia insignis
    auct. non (Lesquereux) Cockerell of Berry, 1929
  • Dipteronia americana
    Brown, 1935 (part)
B. insignis synonymy
  • Myrica insignis
    Lesquereux, 1878
  • Myrica latiloba var. aculiloba Lesquereux, 1878
  • Rhus subrhomboidalis
    Lesquereux, 1883
  • Comptonia acutiloba
    (Lesquereux) Cockerell, 1906
  • Comptonia insignis
    (Lesquereux) Cockerell, 1906
  • Myrica uglowi
    Berry, 1926

Bohlenia is an extinct flowering plant genus in the soapberry family Sapindaceae which is solely known from Eocene sediments exposed in western North America. The genus contains two described species, the type species Bohlenia americana along with Bohlenia insignis. Early description of the genus included both leaves and fruits, however the fruits were removed and named Dipteronia brownii 15 years later. The genus had been considered a part of the tribe Paullinieae, but is currently placed incertae sedis within Sapindaceae.

Distribution

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Bohlenia has been recovered from a series of four geologic formations in the Eocene Okanagan Highlands, with two occurrences of B. americana, one of Cf. B. insignis an one only reported as Bohlenia. Leaves of B. americana are known from the Klondike Mountain Formation sites around Ferry County, Washington, and the Tranquille Formations Falkland site near Cache Creek, British Columbia. The Quilchena site of the Coldwater Beds has undescribed leaves which are only attributed to Bohlenia and not identified to a species.[1][2][3][4][5]

Bohlenia insignis is primarily known from the younger Florissant Formation in Colorado, where it is a notable component of the flora.[2] In contrast, the Chu Chua Formations Joseph Creek site has leaves which were initially synonymized into B. insignis, but have more recently been considered as similar to that species but not part.[2]

Early estimates of the highlands sites ranged from Miocene to Eocene in age. The age of the Klondike Mountain Formation was debated for many years, with plant fossils suggesting a Late Oligocene or Early Miocene age, and the first descriptions of species from the area included them in the Middle Miocene Latah Formation. By the early 1960's the Klondike Mountain formation was thought to be late Oligocene in age.[6] Potassium-argon radiometric dating of samples taken near the Tom Thumb mine in 1966 resulted in a tentative 55 million years old age. Further refinement of sample dating has yielded an approximately Early Eocene, Ypresian age, being radiometrically dated as 49.4 million years old.[3][7][8] A 2003 report using dating of detrital zircon crystals with the tuffs of the Klondike Mountain Formation had been dated to 49.42 ± 0.54 million years ago, the youngest of the Okanagan Highlands sites,[9][10] A 2021 report revised the possible oldest age to around 51.2 ± 0.1 million years ago based on isotopic data from zircon crystals.[11]

The Florissant Formation is composed of successive lake deposits resulting from a volcanic debris flow damming a valley. When Polystoechotites piperatus was described, the Florissant Formation was considered to be Miocene in age, based on the flora and fauna preserved.[12][13] Successive research and fossil descriptions moved the age older and by 1985 the formation had been reassigned to an Oligocene age.[14] Further refinement of the formation's age using radiometric dating of sanidine crystals has resulted in an age of 34.07 million years ago placing the formation in the Priabonian stage of the Late Eocene.[15][16]

The Florissant paleoforest surrounding the lake has been described as similar to modern southeastern North America, with a number of taxa represented that are now found in the subtropics to tropics and confined to the old world. Harry MacGinitie (1953) suggested a warm temperate climate based on the modern biogeographic relatives of the biota found in the formation. Modern estimates of the paleoelevation range between 1,900–4,133 m (6,234–13,560 ft), notably higher than the original estimates by MacGinitie of 300–900 m (980–2,950 ft). Estimates of the mean annual temperature for the Florissant Formation have been derived from climate leaf analysis multivariate program (CLAMP) analysis and modern forest equivalencies of the paleoflora. The results of the various methods have gaven a mean annual temperature rage between approximately 10.8–17.5 °C (51.4–63.5 °F), while the bioclimactic analysis for suggests mean annual precipitation amounts of 50 cm (20 in).[17]

History and classification

[edit]
A Dipteronia brownii samara
Republic, Washigton

The first fossils of Bohlenia were described by Leo Lesquereux (1873) and named Myrica latiloba acutiloba. Two years later (1875) he named Myrica insignis and Myrica saportana, with Rhus subrhomboidalis being named in 1883. While documenting the described Florissant Formation biota as of 1906, Theodore Cockerell transferred both Myrica insignis and M. latiloba acutiloba to Comptonia, but no commentary was made regarding the other two taxa, similarly Frank Knowlton in 1917 reviewed the Florissant specimens and agreed with the placement of the leaf species in Comptonia.[18][19] In 1929 a pair of fossil leaves was described from the Republic Flora in Republic, Washington, which Edward W. Berry then assumed belonged to the Miocene Latah Formation. Berry placed the leaves as Comptonia insignis members and they were retained there until 1935.[20] By 1935 the Republic Flora was recognized to be older than the Latah Formation, though the host Klondike Mountain Formation wouldn't officially be described and named until 1962.[21] Working with new collections from Republic and other sites, in 1935 Roland Wilbur Brown moved the Republic leaves out of Comptonia insignis to a new Dipteronia species Dipteronia americana. He based his expanded description and new species on newly identified fruits from the same location which he deemed the fruits belonging to the leaf fossils.[22] As with the Republic material, further specimen collecting uncovered fruits of Dipteronia present in the Florissant Formation, and so Brown 2 years later united the leaves and fruits. During his re-examination of western United States fossil floras, Brown (1937) redescribed the Florissant material as Dipteronia insignis, with commentary on the variable nature of Dipteronia foliage explaining the synonymizing of Comptonia acutiloba and Rhus subrhomboidalis into the species.[23]

Description

[edit]

The leaves of Bohlenia are thought to have been compound with pinnately formed leaflets. The leaflets have a weak to strongly asymmetrical base and centrally positioned petiole. The margines are serrate and have both simple and compound teeth. Venation of the leaflets is pinnate with a single central main vein running base to apex. Secondary veins branch from the main vein in an alternating pattern, and the major secondaries run to the tooth tips, while minor secondaries fork with branches turning basally and apically. Tertiary veins present between the secondaries run towards the next secondary apically or basally and merge with tertiaries sprouted from that secondary near the mid pint between them.[2][24]

Bohlenia americana

[edit]

[24]

Bohlenia insignis

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Bohlenia insignis holotype USNM-P538

Cf. Bohlenia insignis and undescribed fossils

[edit]

Paleoenvironment

[edit]
Virunga National Park, Albertine Rift, Africa

The Republic and Chu Chua sites are part of a larger fossil site system collectively known as the Eocene Okanagan Highlands. The highlands, including the Early Eocene formations between Driftwood Canyon at the north and Republic at the south, have been described as one of the "Great Canadian Lagerstätten"[25] based on the diversity, quality and unique nature of the paleofloral and paleofaunal 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.[25] The warm temperate highland floras in association with downfaulted lacustrine basins and active volcanism are noted to have no exact modern equivalents. This is due to the more seasonally equitable conditions of the Early Eocene, resulting in much lower seasonal temperature shifts. However, the highlands have been compared to the upland ecological islands of the Virunga Mountains within the African rift valleys Albertine Rift.[26]

The Chu Chua and Klondike Mountain Formations represent a long upland lake system series that was surrounded by a warm temperate ecosystem[27] with nearby volcanism[25] dating from during and just after the early Eocene climatic optimum. The Okanagan Highlands 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.[3] The paleoforest surrounding the lakes have been described as precursors to the 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 range between 0.7 and 1.2 km (0.43 and 0.75 mi) higher than the coastal forests. This is consistent with the paleoelevation estimates for the lake systems, which range between 1.1 and 2.9 km (1,100 and 2,900 m), which is similar to the modern elevation 0.8 km (0.50 mi), but higher.[3]

Estimates of the mean annual temperature have been derived from climate leaf analysis multivariate program (CLAMP) analysis and leaf margin analysis (LMA) of the Republic paleoflora. The CLAMP results after multiple linear regressions gave a mean annual temperature of approximately 8.0 °C (46.4 °F), with the LMA giving 9.2 ± 2.0 °C (48.6 ± 3.6 °F).[3] A bioclimatic-based estimate based on modern relatives of the taxa found at Republic suggested mean annual temperatures around 13.5 ± 2.2 °C (56.3 ± 4.0 °F).[3] This is lower than the mean annual temperature estimates given for the coastal Puget Group, which is estimated to have been between 15 and 18.6 °C (59.0 and 65.5 °F). The bioclimatic analysis for Republic suggests a mean annual precipitation amount of 115 ± 39 cm (45 ± 15 in).[3]

The Florissant paleoforest surrounding the lake has been described as similar to modern southeastern North America, with a number of taxa represented that are now found in the subtropics to tropics and confined to the old world. MacGinitie (1953) suggested a warm temperate climate based on the modern biogeographic relatives of the biota found in the formation. Modern estimates of the paleoelevation range between 1,900–4,133 m (6,234–13,560 ft), notably higher than the original estimates by MacGinitie of 300–900 m (980–2,950 ft). Estimates of the mean annual temperature for the Florissant Formation have been derived from climate leaf analysis multivariate program (CLAMP) analysis and modern forest equivalencies of the paleoflora. The results of the various methods have given a mean annual temperature rage between approximately 10.8–17.5 °C (51.4–63.5 °F), while the bioclimatic analysis for suggests mean annual precipitation amounts of 50 cm (20 in).[17]

References

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  1. ^ Berry, E. (1926). Tertiary floras from British Columbia (PDF) (Report). Geological series; Contributions to Canadian Paleontology. Ottawa, Canada: Geological Survey of Canada. pp. 91–116.
  2. ^ a b c d Wolfe, J.A.; Wehr, W.C. (1987). Middle Eocene dicotyledonous plants from Republic, northeastern Washington (Report). Bulletin. Vol. 1597. United States Geological Survey. pp. 1–25. doi:10.3133/b1597.
  3. ^ a b c d e f g Greenwood, D.; Archibald, S.; Mathewes, R.; Moss, P. (2005). "Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape". Canadian Journal of Earth Sciences. 42 (2): 167–185. Bibcode:2005CaJES..42..167G. doi:10.1139/e04-100. Cite error: The named reference "Greenwood2005" was defined multiple times with different content (see the help page).
  4. ^ Smith, R.Y.; Basinger, J.F.; Greenwood, D.R. (2012). "Early Eocene plant diversity and dynamics in the Falkland flora, Okanagan Highlands, British Columbia, Canada". Palaeobiodiversity and Palaeoenvironments. 92 (3): 309–328. doi:10.1007/s12549-011-0061-5. S2CID 129448108.
  5. ^ 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, U.S.A." Canadian Journal of Earth Sciences. 53 (6): 548–564. doi:10.1139/cjes-2015-0177.
  6. ^ Pearson, R.C. (1967). Geologic map of the Bodie Mountain quadrangle, Ferry and Okanogan counties, Washington (Geologic Quadrangle). Report. United States Geological Survey. doi:10.3133/gq636.
  7. ^ Moss, PT; Greenwood, DR; Archibald, SB (2005). "Regional and local vegetation community dynamics of the Eocene Okanagan Highlands (British Columbia – Washington State) from palynology". Canadian Journal of Earth Sciences. 42 (2): 187–204. Bibcode:2005CaJES..42..187M. doi:10.1139/E04-095.
  8. ^ Archibald, S.B.; Bossert, W.H.; Greenwood, D.R.; Farrell, B.D. (2010). "Seasonality, the latitudinal gradient of diversity, and Eocene insects". Paleobiology. 36 (3): 374–398. Bibcode:2010Pbio...36..374A. doi:10.1666/09021.1. S2CID 55208851.
  9. ^ Archibald, S. B.; Makarkin, V. N. (2021). "Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation". Zootaxa. 4951 (1): 41–79. doi:10.11646/zootaxa.4951.1.2. PMID 33903413. S2CID 233411745.
  10. ^ Archibald, S. B.; Cannings, R. A.; Erickson, R. J.; Bybee, S. M.; Mathewes, R. W. (2021). "The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America". Zootaxa. 4934 (1): zootaxa.4934.1.1. doi:10.11646/zootaxa.4934.1.1. PMID 33756770.
  11. ^ 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. 422: Article 105969. Bibcode:2021SedG..42205969R. doi:10.1016/j.sedgeo.2021.105969. S2CID 237717862.
  12. ^ Cite error: The named reference Cockerell1908 was invoked but never defined (see the help page).
  13. ^ Carpenter, F. M. (1930). "The fossil ants of North America" (PDF). Bulletin of the Museum of Comparative Zoology. 70: 1–66.
  14. ^ Tindale, N. B. (1985). "A butterfly-moth (Lepidoptera:Castniidae) form the Oligocene shales of Florissant, Colorado" (PDF). Journal of Research on the Lepidoptera. 24 (1): 31–40. doi:10.5962/p.266764. S2CID 109301568.
  15. ^ Ksepka, D.T.; Clarke, J.A. (2009). "Affinities of Palaeospiza bella and the Phylogeny and Biogeography of Mousebirds (Coliiformes)". The Auk. 126 (2). The American Ornithologists' Union: 245–259. doi:10.1525/auk.2009.07178. S2CID 85597698.
  16. ^ Lloyd, K.J.; Eberle, J.J. (2008). "A New Talpid from the Late Eocene of North America". Acta Palaeontologica Polonica. 53 (3). Institute of Paleobiology, Polish Academy of Sciences: 539–543. doi:10.4202/app.2008.0311.
  17. ^ a b Smith, D.M. (2008). "A comparison of plant-insect associations in the middle Eocene Green River Formation and the Upper Eocene Florissant Formation and their climatic implications". In Meyer, H. W.; Smith, D. M. (eds.). Paleontology of the Upper Eocene Florissant Formation, Colorado. Special Papers. Vol. 435. Geological Society of America. p. 90. doi:10.1130/2008.2435(06). ISBN 9780813724355.
  18. ^ Knowlton, F.H. (1916). "A review of the fossil plants in the United States National Museum from the Florissant lake beds at Florissant, Colorado, with descriptions of new species and list of type-specimens". Proceedings of the United States National Museum. 51 (2151): 241–297.
  19. ^ LaMotte, R.S. (1952). Catalogue of the Cenozoic plants of North America through 1950. Geological Society of America Memoirs. Vol. 51. Geological Society of America. doi:10.1130/MEM51.
  20. ^ Berry, E.W. (1929). "A revision of the flora of the Latah Formation". USGS Professional Paper Series (PDF). Vol. PP 154-H. pp. 225–265. doi:10.3133/pp154H.
  21. ^ Muessig, Siegfried (1962). "Tertiary volcanic and related rocks of the Republic area, Ferry County, Washington". Geological Survey Research 1962. 450 D: D56–58.
  22. ^ Brown, R. (1935). "Miocene leaves, fruits, and seeds from Idaho, Oregon, and Washington". Journal of Paleontology. 9: 572–587.
  23. ^ Brown, R. W. (1937). Additions to some fossil floras of the Western United States (PDF) (Report). Professional Paper. Vol. 186. United States Geological Survey. pp. 163–206. doi:10.3133/pp186J.
  24. ^ a b Jud, N. A.; Allen, S. E.; Nelson, C. W.; Bastos, C. L.; Chery, J. G. (2021). "Climbing since the early Miocene: The fossil record of Paullinieae (Sapindaceae)". PLOS ONE. 16 (4): e0248369. Bibcode:2021PLoSO..1648369J. doi:10.1371/journal.pone.0248369. PMC 8026063. PMID 33826635.
  25. ^ a b c 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.
  26. ^ DeVore, M. L.; Nyandwi, A.; Eckardt, W.; Bizuru, E.; Mujawamariya, M.; Pigg, K. B. (2020). "Urticaceae leaves with stinging trichomes were already present in latest early Eocene Okanogan Highlands, British Columbia, Canada". American Journal of Botany. 107 (10): 1449–1456. doi:10.1002/ajb2.1548. PMID 33091153. S2CID 225050834.
  27. ^ Cite error: The named reference Pigg2021 was invoked but never defined (see the help page).


† Category:Plants described in 2001 Category:Fossil taxa described in 2001 Category:Eocene plants of North America Category:Fossil record of plants Category:Chu Chua Formation Category:Florissant Formation Category:Klondike Mountain Formation Category:Tranquille Formation Category:Extinct flora of North America

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