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''Nuphar carlquistii'' 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 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
Nymphaeaceae Nymphaeaceae () is a family of flowering plants, commonly called water lilies. They live as rhizomatous aquatic herbs in temperate and tropical climates around the world. The family contains nine genera with about 70 known species. Water li ...
related to the modern spatterdock, ''
Nuphar advena ''Nuphar advena'' (spatterdock or cow lily or yellow pond-lily) is a species of ''Nuphar'' native throughout the eastern United States and in some parts of Canada, such as Nova Scotia.Flora of North America''Nuphar advena''/ref> It is similar to ...
''. 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 ...
seeds and fruits found in the early
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
Okanagan Highlands The Okanagan Highland is an elevated hilly plateau area in British Columbia, Canada, and the U.S. state of Washington (where it is spelled Okanogan Highlands). Rounded mountains with elevations up to above sea level and deep, narrow valleys are ...
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 ...
and
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

''Nuphar carlquistii'' fossils have been identified from two locations in Western North America, the 49 million year old
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 ...
near
Republic, Washington Republic is a city in Ferry County, Washington, United States. The population was 1,073 at the 2010 census, a 12.5% increase over the 2000 census. It is the county seat of Ferry County. It was the largest mining camp in the Republic Mining Distr ...
and at the Thomas Ranch locality 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 ...
. Permineralized seeds identified as the extinct Nymphaeaceae genus '' Allenbya'' has been identified from the related Okanagan Highlands
Princeton chert The Princeton Chert is a fossil locality in British Columbia, Canada, which comprises an anatomically preserved flora of Eocene Epoch age, with rich species abundance and diversity. It is located in exposures of the Allenby Formation on the east ...
site in south central British Columbia. Ages for the Okanagan Highland locations are, in general,
Early Eocene 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 i ...
, 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 (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
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 Midd ...
in age. The 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 Melanie DeVore, Witt Taylor, and Kathleen Pigg 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 ''N. carlquistii'' being published in a 2015 ''
International Journal of Plant Sciences The ''International Journal of Plant Sciences'' covers botanical research including genetics and genomics, developmental and cell biology, biochemistry and physiology, morphology and structure, systematics, plant-microbe interactions, paleobotany, ...
'' article. They chose the
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''carlquistii'', in honor of
Sherwin Carlquist Sherwin John Carlquist FMLS (July 7, 1930 - December 1, 2021) was an American botanist and photographer. Education He received his undergraduate degree from the University of California, Berkeley in 1952 and a Ph.D. in botany in 1956, also at ...
in recognition of the work he had done with Nymphaeaceae anatomy and with plant biology. In the type description, the species name was applied to fruits and seeds specifically, while fossils of associated but not attached
tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s, stigmatic disks and
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
sections were discussed as ''Nuphar'' cf ''carlquistii''. One of the rhizome segments, UWBM 57225A showing root attachment points, had previously been published with an informal identification as banana fruit of the genus ''
Ensete ''Ensete'' is a genus of monocarpic flowering plants native to tropical regions of Africa and Asia. It is one of the three genera in the banana family, Musaceae, and includes the false banana or enset ('' E. ventricosum''), an economically impo ...
''. The specimen was figured in a 1996 article with a caption of "''Ensete'' sp. fruits with seeds, UWBM 57225A, loc. A0307" though no discussion of the fossil was made in the article text. The fossils display a combination of features, such as a barrel shaped fruit that has vertical furrows, with the stigmatic disk being occasionally to fully crenulated, that are seen in ''Nuphar'' Section ''Astylus'', and are noted to be quite similar to the living ''Nuphar advena''.


Description

The fruits are between and across, expanding up from a base. The fruits have a grouping of five persistent tepals that surround the fruit and which have groupings of
sclereid Sclereids are a reduced form of sclerenchyma cells with highly thickened, lignified cellular walls that form small bundles of durable layers of tissue in most plants.Evert, Ray F; Eichhorn, Susan E. Esau's Plant Anatomy: Meristems, Cells, and Ti ...
s that vary in size. One specimen has at least sixteen
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s that were preserved compressed against the tepals, while another shows approximately ten seeds in growth position in the fruit. Seeds are also found preserved individually and as groupings or masses of seeds ranging between and . One of the specimens has several hundred seeds grouped closely together and enclosed in a thick matrix, thought to be the preserved jelly like matrix that the seeds first dispersed from the fruit in. Other masses show lesser amounts of matrix, and the ones showing little to no matrix are less grouped together. The individual seeds range between long with an oval to barrel shape and an operculum at one end. Each seed has a distinct
raphe Raphe (; from Greek ῥαφή, "seam"Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with the assistance of. Roderick McKenzie.'' Oxford: Clarendon Press.) has several diffe ...
ridge running vertically down it from the base to the slightly asymmetrical operculum. While most of the seed coat structure is not preserved enough to identify, the outer cell layer is distinct, showing small pentagon shaped cells with straight walls, a feature seen in fossil and living ''Nuphar''.


References


External links

* {{Taxonbar, from=Q29025327 Nymphaeaceae Ypresian plants of North America Extinct flora of North America Paleontology in Washington (state) Paleogene British Columbia Paleontology in British Columbia Klondike Mountain Formation Fossil taxa described in 2015