Ulmus okanaganensis
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''Ulmus okanaganensis'' is an extinct species of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
in the family
Ulmaceae The Ulmaceae () are a family of flowering plants that includes the elms (genus ''Ulmus''), and the zelkovas (genus ''Zelkova''). Members of the family are widely distributed throughout the north temperate zone, and have a scattered distribution ...
related to the modern
elm Elms are deciduous and semi-deciduous trees comprising the flowering plant genus ''Ulmus'' in the plant family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical-montane regions of North ...
s. The species is known from
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 ...
leaves, flowers, and fruits found in the early
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
deposits of northern
Washington state Washington (), officially the State of Washington, is a state in the Pacific Northwest region of the Western United States. Named for George Washington—the first U.S. president—the state was formed from the western part of the Washington ...
, United States and similar aged formations in
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, ...
, Canada.


History and classification

''U. okanaganensis'' fossils have been identified from five locations in Western North America. The type description listed occurrences at the 49 million year old Klondike Mountain Formation near Republic, Washington plus the British Columbian sites of One Mile Creek near
Princeton, British Columbia Princeton (originally Vermilion Forks) is a town in the Similkameen region of southern British Columbia, Canada."The rich history of Princeton or how Vermilion Forks made it on the map...", Princeton 2008 Visitors Guide, p. 4. It lies just east o ...
, the
McAbee Fossil Beds The McAbee Fossil Beds is a Heritage Site that protects an Eocene Epoch fossil locality east of Cache Creek, British Columbia, Canada, just north of and visible from Provincial Highway 97 / the Trans-Canada Highway ( Highway 1). The McAbee Fo ...
east of Cache Creek, and in the
Driftwood Canyon Provincial Park Driftwood Canyon Provincial Park is a provincial park in British Columbia, Canada. Driftwood Canyon Provincial Park covers 23 ha of the Bulkley River Valley, on the east side of Driftwood Creek, a tributary of the Bulkley River, 10 km northe ...
northeast of
Smithers Smithers is a surname of English origin. It derives from the Middle English term "smyther", referring to a metalsmith, and is thus related to the common occupational surname Smith. The name Smither is related. People *Alan Smithers (born 1938 ...
. In 2009 the species was reported from a fourth British Columbian site near Falkland. Ages for the Okanagan Highland locations are, in general, Early Eocene, with the sites that have current uranium-lead or argon–argon radiometric dates being of
Ypresian In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian ...
age, while the undated sites or those given older dates being possibly slightly younger and
Lutetian The Lutetian is, in the geologic timescale, a stage or age in the Eocene. It spans the time between . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it is sometimes referred to as the ...
in age. Roland Brown examined fossils from the Klondike Mountain formation in a 1936 paper and placed some leaves into the extinct species '' Zelkova oregoniana'' along with fossils from the
Latah Formation The Latah Formation is a series of late Miocene lacustrine sedimentary deposits which outcrop in eastern Washington and northwestern Idaho. The lake beds are interbedded with igneous rock of the Columbia River Basalt Group. The formation was ...
in Washington, which the Republic fossils were thought to belong to, and 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 ...
in Oregon. Ulmaceous fruits from the Princeton area had been tentatively identified as a species of '' Chaetoptelea'', a southern North American genus that is now included into ''Ulmus''. The discovery of leaves attached to branchlets with fruits, and attached to branchlets attached to flowers at several fossil sites led to a restudy of the Okanagan highlands leaves and fruits. Fossils of leaves and fruits that had previously been identified as ''Zelkova'' and ''Chaetoptelea'' along with additional specimens were studied by
paleobotanist Paleobotany, which is also spelled as palaeobotany, is the branch of botany dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeogr ...
s Thomas Denk and Richard Dillhoff, with the
type description A species description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have be ...
for ''U. okanaganensis'' being published in a 2005 ''
Canadian Journal of Botany ''Botany'' is a monthly peer-reviewed scientific journal that is published by Canadian Science Publishing. It was established in 1935 as the ''Canadian Journal of Research, Section C: Botanical Sciences'' and renamed in 1951 as ''Canadian Journal ...
'' article. They chose the specific name ''okanaganensis'', in reference to the Okanagan Highlands, of Central British Columbia and northern Washington, where the species is a common paleofloral element of the Eocene lake deposits in the region. Based on phylogenetic analysis that was performed by Denk and Dillhoff, ''U. okanaganensis'' was placed as a member of ''Ulmus'' subgenus ''Ulmus'', between ''U.'' section ''Microptelea'' and ''U.'' section ''Ulmus''. Denk and Dillhoff noted that the results of the analysis also suggested the subgenus to be a
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basal to the derived species in ''Ulmus'' subgenus ''Oreoptelea''. The morphology of the leaf edges and bases was noted to be similar to fossils found in Paleocene rocks of Ellesmere Island, late Paleocene to Early Eocene rocks of Spitsbergen, and Paleocene rocks of North-east China. The species Early Eocene '' Ulmus fushunensis'' described in 2010 from the Jijuntun Formation of Liaoning Province, China, displays noted similarities to ''U. okanaganensis'' as well.


Description

The studied leaves of ''U. okanaganensis'' are split into three major morphotypes, leaves from sucker-shoot stems of the trees, leaves from the "elongation" or short branches, and leaves from the reproductive shoots. The sucker-shoot leaves range up to in length. The short, thick petiole is bracketed by uneven lobes of the leaf base, with one lobe typically being distinctly larger than the other. The leaves have a pinnate vein structure, with the primary vein having between twelve and fourteen secondary veins branching from it at decreasingly sharp angles from apex to base of the leaf. The secondary veins and the exterior branches off the secondaries terminate in teeth along the leaf margins. The leaves have compound teeth, each of the large teeth on having a smaller tooth located basally between it and the next large tooth. All the teeth typically have convex sides that terminate at blunt tooth tips, though occasional teeth have straight or concave sides. In the elongation shoot leaves, the petioles are short and thick, though shorter and thinner than those of the sucker-shoot leaves. Similarly the leaves are smaller, ranging up to long, and the bases are typically only slightly asymmetrical. They have a
cordate Cordate is an adjective meaning ' heart-shaped' and is most typically used for: * Cordate (leaf shape) The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a ...
to auriculate leaf blade base and an acute leaf tip. The primary vein ranges between fully straight and slightly curved as it progresses from base to tip. There are a similar number of secondary veins to those in the other two leaf types, with a usual minimum of eleven pairs; however, the angle of the secondaries to the primary increases towards the base rather than decreasing as seen in the sucker-shoot leaves. The short-shoot leaves are range between and long, much like the length of the elongation shoot leaves. However the elongation shoot leaves are up to wide while short shoot leaves are wide. There are between eight and fourteen secondary veins, with up to three veins branching off the undersides of each secondary close to the leaf base, though one to no veins branch from the secondaries near the leaf tip. The teeth on short-shoot leaves are simple, with the apical side of each tooth usually concave and the basal side usually convex. The flowers and fruits develop on fascicles at the leaf nodes of branches. The fruits are small, ranging between , with a
pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
up to long. Like species of the modern ''Ulmus'' section ''Chaetoptelea'', the fruits do not have a surrounding wing, and the styles on the fruit apex are free. The ''U.'' sect. ''Chaetoptelea'' fruits are noted to be distinctly ciliate, having many fine hairs on the fruits, but only one fossil has been described showing ciliate, the other specimens are all smooth, possibly as a result of processes before fossilization.


References

{{Taxonbar, from=Q21396866 †okanaganensis Ypresian plants of North America Extinct flora of North America Paleontology in Washington (state) Paleontology in British Columbia Fossil taxa described in 2005 Allenby Formation Klondike Mountain Formation Chu Chua Formation Coldwater Beds Tranquille Formation