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''Dipteronia brownii'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
species in the soapberry family (Sapindaceae) described in 2001.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s of ''D. brownii'' are known from stratigraphic formations in North America and Asia ranging in age between
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
to
Early Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chatti ...
.


Distribution and paleoecology

The oldest occurrences for ''Dipteronia brownii'' are both Paleogene in age, with fossils found in the Middle Paleocene,
Fort Union Formation The Fort Union Formation is a geologic unit containing sandstones, shales, and coal beds in Wyoming, Montana, and parts of adjacent states. In the Powder River Basin, it contains important economic deposits of coal, uranium, and coalbed methane. ...
of Wyoming and the
Danian The Danian is the oldest age or lowest stage of the Paleocene Epoch or Series, of the Paleogene Period or System, and of the Cenozoic Era or Erathem. The beginning of the Danian (and the end of the preceding Maastrichtian) is at the Cretaceous†...
Middle-Upper Tsagayan Formation of Northeastern coastal Russia. Migration between the slightly older Russian Far East site and North America was likely facilitated by the Beringian land bridge during the early to middle Paleoene.. In the Early Eocene the species expanded into the
Eocene Okanagan Highlands The Eocene Okanagan Highlands or Eocene Okanogan Highlands are a series of Early Eocene geological formations which span a transect of British Columbia, Canada, and Washington state, United States. Known for a highly diverse and detailed Paleoflo ...
sites of East central
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
and north east central
Washington Washington commonly refers to: * Washington (state), United States * Washington, D.C., the capital of the United States ** A metonym for the federal government of the United States ** Washington metropolitan area, the metropolitan area centered o ...
. Fossils have been found in the Okanagan highlands formations from the southern most
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington state. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Wash ...
to the northern most Driftwood Shales, with occurrences in the
Allenby Formation 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 abun ...
, Tranquille Formation,
Chu Chua Formation Chu or CHU may refer to: Chinese history * Chu (state) (c. 1030 BC–223 BC), a state during the Zhou dynasty * Western Chu (206 BC–202 BC), a state founded and ruled by Xiang Yu * Chu Kingdom (Han dynasty) (201 BC–70 AD), a kingdom of the H ...
and
Horsefly Shales Horse-flies or horseflies are true flies in the family Tabanidae in the insect order Diptera. They are often large and agile in flight, and only the female horseflies bite animals, including humans, to obtain blood. They prefer to fly in su ...
. During the Middle Eocene the species appears in the
John Day Formation The John Day Formation is a series of rock strata exposed in the Picture Gorge district of the John Day River basin and elsewhere in north-central Oregon in the United States. The Picture Gorge exposure lies east of the Blue Mountain uplift, whi ...
of central Oregon. During the middle to late Eocene the species spread east and south to the Ruby Basin Flora of Montana and the
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones and ...
of Colorado, while the last North American occurrence is in the Early Oligocene, Rupelian of upper John Day Formations Bridge Creek Flora. Concurrently, ''Dipteronia brownii'' fruits are also found in
Rupelian The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...
lacustrine A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much larger ...
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
s of
Chuxiong Yi Autonomous Prefecture Chuxiong Yi Autonomous Prefecture (; Chuxiong Yi script: ,IPA: ; Yi script: ꊉꇑꆑꌠꑼꂰ; Yi Pinyin: wop lup nut su yuop mi) is an autonomous prefecture located in central Yunnan Province, China. Chuxiong has an area of . The capital of t ...
southwestern
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, and are also the southern most of the ''D. brownii'' fossils. Ding ''et al.'' (2018) posited that the reduction of ''Dipteronia'' from the broad North American and wide Asian range seen in ''D. brownii'' to the isolated regional
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
of modern times seen in '' Dipteronia dyeriana'' and '' Dipteronia sinensis'' was the result of several factors. Range reduction was likely due to overall global cooling during the Oligocene and Miocene, combined with increased and intensified rain fall in the northern hemisphere and associated high latitude drying.


History and classification

Roland Brown (1935) described the ''Dipteronia'' species ''Dipteronia americana'' based on both fruits and leaflets including one pair of leaflets first figured by
Edward Berry Rear Admiral Sir Edward Berry, 1st Baronet, KCB (17 April 1768 – 13 February 1831) was an officer in Britain's Royal Navy primarily known for his role as flag captain of Rear Admiral Horatio Nelson's ship HMS ''Vanguard'' at the Battle of ...
(1929). The species was redescribed by Jack Wolfe and
Wesley Wehr Wesley Conrad Wehr (April 17, 1929 – April 12, 2004) was an American paleontologist and artist best known for his studies of Cenozoic fossil floras in western North America, the Stonerose Interpretive Center, and as a part of the Northwest ...
(1987) who designated the leaflet as
lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
for the species, which they moved to the new extinct genus '' Bohlenia'' based on the difference in leaflet venation from that of other sapindalian taxa. Manchester (1999) figured a single ''Dipteronia'' mericarp fossil which was at that time identified as being from the Eocene Fushun Formation in Liaoning Province, China. The location was taken from the specimen drawer in the
University of California Museum of Paleontology The University of California Museum of Paleontology (UCMP) is a paleontology museum located on the campus of the University of California, Berkeley. The museum is within the Valley Life Sciences Building (VLSB), designed by George W. Kelham and ...
collections where the fossil was stored; however, the fossil itself did not have a label indicating its location. Doubt was later raised regarding the provenance of the fossil, with Manchester being informed that the UCMP formerly housed collections of very similarly colored shale from the Chu Chua Formation near Joseph Creek, British Columbia. The Chu Chua fossils had been transferred to the
Geological Survey of Canada The Geological Survey of Canada (GSC; french: Commission géologique du Canada (CGC)) is a Canadian federal government agency responsible for performing geological surveys of the country, developing Canada's natural resources and protecting the en ...
collections in the 1970's, but it was possible the ''Dipteronia'' specimen was left behind.
Palynological Palynology is the "study of dust" (from grc-gre, παλύνω, palynō, "strew, sprinkle" and ''-logy'') or of "particles that are strewn". A classic palynologist analyses particulate samples collected from the air, from water, or from deposit ...
analysis of pollen in the specimen matrix was performed in October 2000 in conjunction with the Manchester (2001) research as an attempt to determine the origin site. Based on
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
analysis of Fushun shale, Chu Chua shale and the specimen, much of the palynofloras overlapped. However the palynomorphs '' Liquidambarpollenites'' and '' Ephedripites'' were only seen in Fushun samples and not the Chu Chua or the mystery sample. Additionally though no other ''Dipteronia'' have been identified in Joseph Creek collections, the genus is present at coeval locations elsewhere in the Eocene Okanagan Highlands, leading Manchester (2001) to place the fossil as Joseph Creek, and state ''Dipteronia'' was not present in the Fushun Flora. The North American ''Dipteronia'' fossils were reexamined and redescribed by Amy McClain and Steven Manchester, whose 2001 type description for ''Dipteronia brownii'' was published in the ''
American Journal of Botany The ''American Journal of Botany'' is a monthly peer-reviewed scientific journal which covers all aspects of plant biology. It has been published by the Botanical Society of America since 1914. The journal has an impact factor of 3.038, as of 201 ...
''. McClain and Manchester noted the lack of attachment fossils uniting the ''Bohlenia americana'' leaflets to fruits, and as such opted to remove the fruits from ''Bohlenia'' and restrict the taxon definition to only foliage. They opted to name the new fruit species ''D. brownii'' as a
patronym A patronymic, or patronym, is a component of a personal name based on the given name of one's father, grandfather (avonymic), or an earlier male ancestor. Patronymics are still in use, including mandatory use, in many countries worldwide, alt ...
honoring Roland Brown as the first systematist to recognize ''Dipteronia'' fossils.


Description

''Dipteronia brownii'' fruits were born in
schizocarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more seeds (the m ...
comprised of three, and less commonly two, flat
mericarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more seeds (the m ...
s which attach along a straight proximal edge. The schizocarps were born on long thin
pedicels In botany, a pedicel is a stem that attaches a single flower to the inflorescence. Such inflorescences are described as ''pedicellate''. Description Pedicel refers to a structure connecting a single flower to its inflorescence. In the absenc ...
which flared to a disk-shaped juncture with the
perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...
. The mericarps have a circular smooth outline, extending from the proximal attachment scar, giving a subelliptical profile to the wing. A single primary vein runs from the pedicel up the attachment scar before turning inward at a 90–135° angle towards the center of the wing. In the central area of the wing, from the wing edge, is a pyriform to elliptical seed with a diameter of . Covering the surface of the seed
pericarp Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Aggre ...
is a
reticulated Reticulation is a net-like pattern, arrangement, or structure. Reticulation or Reticulated may refer to: * Reticulation (single-access key), a structure of an identification tree, where there are several possible routes to a correct identificatio ...
network of secondary veins. The tertiary veins spreading out from the seed though the wing split and join as they extend to the wing margin. connecting the tertiaries is a fine reticulum of quaternary veins which form polygonal areoles.


References

{{Taxonbar, from=Q21399733 † Plants described in 2001 Fossil taxa described in 2001 Eocene plants of North America Fossil record of plants Allenby Formation Coldwater Beds Florissant Formation Horsefly Shales Klondike Mountain Formation Tranquille Formation Chu Chua Formation