''Nyssa spatulata'' is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

in the

tupelo Tupelo commonly refers to: * Tupelo (tree), a small genus of deciduous trees with alternate, simple leaves * Tupelo, Mississippi, the county seat and the largest city of Lee County, Mississippi Tupelo may also refer to: Places * Tupelo, Arka ...

family,

Nyssaceae Nyssaceae is a family of flowering trees sometimes included in the dogwood family (Cornaceae). Nyssaceae is composed of 37 known species in the following five genera:Averyanov, L. V. & Hiep, N. T. (2002). ''Diplopanax vietnamensis'', a New Speci ...

known from the

middle Eocene The Eocene ( ) is a geological epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''Ēṓs'', ' Dawn') a ...

sediments exposed in north central

Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...

. The species was first described from a series of isolated

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 preserve ...

seeds in

chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a prec ...

History and classification

''N. spatulata'' has been identified from a single location in the

Clarno Formation John Day Fossil Beds National Monument is a National monument (United States), U.S. national monument in Wheeler County, Oregon, Wheeler and Grant County, Oregon, Grant counties in east-central Oregon. Located within the John Day River basin an ...

, the Clarno nut beds, type locality for both the formation and the species. The nut beds are approximately east of the unincorporated community of

Clarno, Oregon Clarno is an unincorporated community in Wasco County, Oregon, United States. It is located along Oregon Route 218 near the John Day River. History The community was named after Andrew Clarno, an early settler. The Clarno post office was est ...

, and considered to be middle Eocene in age, based on averaging zircon fission track

radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to Chronological dating, date materials such as Rock (geology), rocks or carbon, in which trace radioactive impurity, impurities were selectively incorporat ...

which yielded an age of 43.6 and 43.7 ± 10

million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...

and

Argon–argon dating Argon–argon (or 40Ar/39Ar) dating is a radiometric dating method invented to supersede Potassium-argon dating, potassiumargon (K/Ar) dating in accuracy. The older method required splitting samples into two for separate potassium and argon measur ...

which yielded a 36.38 ± 1.31 to 46.8 ± 3.36 mya date. The average of the dates resulted in an age range of 45 to 43 mya. The beds are composed of silica and calcium carbonate cemented

tuff Tuff is a type of rock made of volcanic ash ejected from a vent during a volcanic eruption. Following ejection and deposition, the ash is lithified into a solid rock. Rock that contains greater than 75% ash is considered tuff, while rock co ...

aceous sandstones, siltstones, and conglomerates which preserve either a lake delta environment, or alternatively periodic floods and volcanic mudflows preserved with hot spring activity. The species was described from a series of

type specimens 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 associated. In other words, a type is an example that serves to anchor or centralizes t ...

, the

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

specimen USNM 422378, which was preserved in the paleobotanical collections of the

National Museum of Natural History The National Museum of Natural History (NMNH) is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. With 4.4 ...

Washington, D.C. Washington, D.C., formally the District of Columbia and commonly known as Washington or D.C., is the capital city and federal district of the United States. The city is on the Potomac River, across from Virginia, and shares land borders with ...

and four

paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...

specimens. Two of the paratypes were also in the national Museum collections, while the remaining two were in the

University of Florida The University of Florida (Florida or UF) is a public university, public land-grant university, land-grant research university in Gainesville, Florida, United States. It is a senior member of the State University System of Florida and a preem ...

collections in

Gainesville, Florida Gainesville is the county seat of Alachua County, Florida, United States, and the most populous city in North Central Florida, with a population of 145,212 in 2022. It is the principal city of the Gainesville metropolitan area, Florida, Gainesv ...

. The fossils were part of a group of approximately 20,000 specimens collected from 1942 to 1989 by Thomas Bones, Alonzo W. Hancock, R. A. Scott, Steven R. Manchester, and a number of high school students. The ''N. spatulata'' specimens were first studied by graduate student R. A. Scott, who placed the species into the extinct genus ''Palaeonyssa'' as ''Palaeonyssa spatulata'', with the 1954

type description A species description is a formal scientific description of a newly encountered species, typically articulated through a scientific publication. Its purpose is to provide a clear description of a new species of organism and explain how it differ ...

of the species appearing in the journal ''

Palaeontographica This is a list of academic journals, monographic series, and other serials published by E. Schweizerbart. A B C D E F G H I J K L M N P Q R S T U V W Z {{columns-list, colwidth=30em, *''Zeitschrift ...

''. ''Palaeonyssa'' was first described by Eleanor Reid and Marjorie Chandler from fossils preserved in the

London Clay The London Clay Formation is a Sediment#Shores and shallow seas, marine formation (geology), geological formation of Ypresian (early Eocene Epoch, c. 54-50 million years ago) age which outcrop, crops out in the southeast of England. The London C ...

, with them describing several species in the genus. ''N. spatulata'' was re-examined by

paleobotanist Paleobotany or palaeobotany, also known as paleophytology, is the branch of botany dealing with the recovery and identification of plant fossils from geological contexts, and their use for the biological reconstruction of past environments (pale ...

Steven R. Manchester of the University of Florida, who published a 1994 re-description for the species in the Journal ''

Palaeontographica Americana The Paleontological Research Institution, or PRI, is a paleontological organization in Ithaca, New York, with a mission including both research and education. PRI is affiliated with Cornell University, houses one of the largest fossil collecti ...

''. In the re-description Manchester noted there was very little difference between the extinct genus ''Palaeonyssa'' and the modern ''Nyssa'', and deemed the two to be the same genus. The merging resulted in the recombination of ''Palaeonyssa spatulata'' as ''Nyssa spatulata''.

Description

The fruits of ''N. spatulata'' have a generally rounded triangular to round

endocarp 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. Fruitlike structures may develop directly from the seed itself rather th ...

in cross-section and are trilocular, similar to the extant '' N. talamancana''. The outline of the endocarps is oblong to ellipsoid, having both apex and base rounded and showing 9 ridges running the length of the endocarp from base to apex. The fruits have an average length of though lengths of are seen. Similarly while the average girth of the fruits is , thicker fruits reaching are known. Each of the three locules has a slit like opening along the upper half of the endocarp, and are separated from each other by a wall of swirling groups of fibers. There is one seed per locule, with the locule having a c-shaped cross section, the chamber opening near the apex and narrowing near the base.

References

{{Taxonbar, from=Q21397408 Eocene plants Extinct flora of North America

† A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...

Fossil taxa described in 1954 spatulata Plants described in 1954 Clarno Formation