''Abies milleri'', 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

fir Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...

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 ...

remains found in deposits from 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' ...

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 ...

stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...

(around 49.5 mya) in

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 on ...

State, USA, is the oldest confirmed record for the

genus.

History and classification

The species was described from 81 fossil specimens collected from

Burke Museum The Burke Museum of Natural History and Culture (Burke Museum) is a natural history museum in Seattle, Washington, in the United States. Established in 1899 as the Washington State Museum, it traces its origins to a high school naturalist club fo ...

site number A0307 in

Ferry County, Washington Ferry County is a county located on the northern border of the U.S. state of Washington. As of the 2020 census, the population was 7,178, making it the fourth-least populous county in Washington. The county seat and largest city is Republic. Th ...

. The

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

specimen, number # ''UWBM 31299'', and the eleven

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 nor a syntype). Of ...

specimens are currently deposited in the collections of the Burke Museum in

Seattle Seattle ( ) is a seaport city on the West Coast of the United States. It is the seat of King County, Washington. With a 2020 population of 737,015, it is the largest city in both the state of Washington and the Pacific Northwest regio ...

, where they were studied and described by Howard E. Schorn and Wesley C. Wehr. Schorn and Wehr published their 1986

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 ''A. milleri'' in the '' Burke Museum Contributions in Anthropology and Natural History'', Volume 1. 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 ...

, ''milleri'', was coined in honor of Charles N. Miller Jr for his contributions to the study and understanding of the conifer family

Pinaceae The Pinaceae, or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as Cedrus, cedars, firs, Tsuga, hemlocks, larches, pines and spruces. The family is included in the order Pinales, ...

. The studied specimens were excavated from the Tom Thumb Tuff

member Member may refer to: * Military jury, referred to as "Members" in military jargon * Element (mathematics), an object that belongs to a mathematical set * In object-oriented programming, a member of a class ** Field (computer science), entries in ...

of the

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 ...

in the city of

Republic A republic () is a "state in which power rests with the people or their representatives; specifically a state without a monarchy" and also a "government, or system of government, of such a state." Previously, especially in the 17th and 18th c ...

. ''A. milleri'' has been placed in the

genus ''

Abies Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family Pinaceae. They are found on mountains throughout much of North and Central America, Europe, Asia, and North Africa. The genus is most closely related to ...

'', which has between 48 and 55 species native to much of North and Central

America The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

and

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

occurring in

mountains A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually higher th ...

over most of the range. The characters used to separate

extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...

species of ''Abies'' are based on the reproductive structures such as cone scales, wing seeds and bracts. The placement in ''Abies'' is based on the dorso-ventrally flattened leaves, and the circular leaf scars, which separate the organs from those of ''

Pseudotsuga ''Pseudotsuga'' is a genus of evergreen coniferous trees in the family Pinaceae (subfamily Laricoideae). Common names for species in the genus include Douglas fir, Douglas-fir, Douglas tree, Oregon pine and Bigcone spruce. '' Pseudotsuga menz ...

'' and ''

Keteleeria ''Keteleeria'' is a genus of three species of coniferous trees in the family Pinaceae first described as a genus in 1866. The genus name ''Keteleeria'' honours J.B. Keteleer (1813–1903), a French nurseryman. The group is related to the genera ...

''. Within ''Abies'', ''A. milleri'' shows similarities with '' A. kawakamii'' and '' A. chensiensis'' from Asia and with '' A. concolor'' and '' A. lasiocarpa'' of North America. ''A. milleri'' does not show traits which allow placement in any one of the genus ''Abies'' sections, however. The 81 specimens studied for the ''A. milleri'' description included 40 cone scales, 21 wing seeds, 10 foliated axes, and two detached needles. Schorn and Wehr placed the different detached organ fossils in the same species based on a number of factors. Where visible, all the cone scales have bracts and are morphologically similar. The scales display impressions on the admedial surface of wing seeds that match the fossil seed dimensions. The needle arrangement is consistent on both foliated and defoliated axes, with attached needles matching isolated needles. All of the fossils occurred in the same stratigraphic layers.

Description

The cone scales are up to and 1.25-2 times as long as wide, have a generally triangular to cuneate shape with the distal edge turned upwards and the stalk on half the length of the scale. Sometimes preserved with cone scales are the bracts which average and have a central accuminate flanked by thinner laminae. The wing seeds are up to and have a pterostegium which covers on average three-quarters of the seed. The narrow obovate seed is generally , 1-1.5 times the length of the upper wing edge while the wing itself is widest in the distal half. Needles of ''A. milleri'' reach up to long, but have a width of only at the base. The leaf base is generally as wide to slightly wider than the leaf and round, with the leaf angled approximately 70 degrees upward from the attachment. Leaf scars on axes are circular to slightly oval, , and show a vascular bundle scar in the center. Oval leaf scars are oriented with the long axis parallel to the length of the stem axis. While the scars show the leaves to be in a helical arrangement, the leaves would have had a flattened orientation on either side of the axis.

References

{{Taxonbar, from=Q2821614 milleri Plants described in 1986 Ypresian plants of North America Extinct flora of North America Klondike Mountain Formation