Paraconcavistylon
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''Paraconcavistylon'' 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 ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
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
Trochodendraceae Trochodendraceae is the only family of flowering plants in the order Trochodendrales. It comprises two extant genera, each with a single species along with up to five additional extinct genera and a number of extinct species. The living specie ...
comprises a single species, ''Paraconcavistylon wehrii''. The genus 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 ...
fruits and leaves 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' ...
deposits of northern Washington state, United States, and southern British Columbia, Canada. The species was initially described as a member of the related extinct genus '' Concavistylon'' as ''"Concavistylon" wehrii'', but subsequently moved to the new genus ''Paraconcavistylon'' in 2020 after additional study.


Distribution and paleoenvironment

''Paraconcavistylon wehrii'' is known from specimens which were recovered from outcrops of the early 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 ...
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
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 ...
and coeval
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 ...
near Cache Creek, British Columbia. The Klondike Mountain Formation and McAbee Fossil sites preserve upland temperate floras which were first interpreted as being
microthermal In climatology, the term microthermal is used to denote the continental climates of Eurasia and North America. Etymology The word ''microthermal'' is derived from two Greek words meaning "small" and "heat". This is misleading, however, since the t ...
, however further study has shown the floras to be more
mesothermal In climatology, the term mesothermal is used to refer to certain forms of climate found typically in the Earth's temperate zones. It has a moderate span of temperature, with winters not cold enough to sustain snow cover. Summers are warm within oc ...
in nature. The plant community preserved in the Klondike Mountain formation is a mixed conifer–broadleaf forest with large pollen elements of
birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech-oak family Fagaceae. The genus ''Betula'' contains 30 ...
and
golden larch ''Pseudolarix amabilis'' is a species of coniferous tree in the pine family Pinaceae. The species is commonly known as golden larch, but being more closely related to ''Keteleeria'', ''Abies'' and ''Cedrus'', is not a true larch (''Larix''). ' ...
, but also having notable traces of fir,
spruce A spruce is a tree of the genus ''Picea'' (), a genus of about 35 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal (taiga) regions of the Earth. ''Picea'' is the sole genus in the subfami ...
,
cypress Cypress is a common name for various coniferous trees or shrubs of northern temperate regions that belong to the family Cupressaceae. The word ''cypress'' is derived from Old French ''cipres'', which was imported from Latin ''cypressus'', the ...
, and
palm Palm most commonly refers to: * Palm of the hand, the central region of the front of the hand * Palm plants, of family Arecaceae **List of Arecaceae genera * Several other plants known as "palm" Palm or Palms may also refer to: Music * Palm (ba ...
.


Taxonomy and phylogeny

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 ...
was originally collected by Dwyane Day 1999 and subsequently donated to the University of Washington's
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 ...
as specimen UWBM PB 101336. One of the paratypes, specimen USNM 537360 is the earliest collected specimen, being recovered in 1901 by then Washington State Geologist
Harry Landis Harry Landis (born Hyman Jacob Londinsky, 25 November 1926 – 11 September 2022) was a British actor and director. He had a long career in theatre, film and television, spanning over 60 years. Landis was best known for playing barber Felix K ...
. Study of the fossil by paleobotanists Steven Manchester ''et al'' resulted in the description of ''"Concavistylon" wehrii'' being published in 2018, 117 years after the Landis specimen was found. The specific name ''wehrii'' was chosen as a patronym honoring Wesley "Wes" Wehr for his enthusiasm and generosity. Often plant fossils are isolated parts such as leaves or seeds, which makes it difficult to identify relationships to other isolated parts. Such fossils from the same species are described as
morphospecies In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. "
Rosetta Stone The Rosetta Stone is a stele composed of granodiorite inscribed with three versions of a Rosetta Stone decree, decree issued in Memphis, Egypt, in 196 BC during the Ptolemaic dynasty on behalf of King Ptolemy V Epiphanes. The top and middle te ...
" fossils are rare cases of multiple portions of a fossil preserved in connection. The holotype of ''Paraconcavistylon'' has been described as a Rosetta Stone fossil as it bears an
infructescence Infructescence (fruiting head) is defined as the ensemble of fruits derived from the ovaries of an inflorescence. It usually retains the size and structure of the inflorescence. In some cases, infructescences are similar in appearance to simple fru ...
attached to branchlet that also has leaves and terminal buds connected, allowing for a fuller plant description than is usually possible. The description of
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 ...
trochodendraceous fossils from Wyoming and a phylogenetic analysis of two living and four extinct genera indicated that ''Concavistylon'' was not monophyletic. Based on the pendulous nature of ''"C." wehrii'' inflorescences, which are distinct from the erect inflorescences of ''C. kvacekii'', the new genus ''Paraconcavistylon'' was erected with ''"C." wehrii'' as the type species. ''Paraconcavistylon wehrii'' is one of between three and four trochodendraceae species that have been described from the Klondike Mountain Formation. Broadly circumscribed three other species have been identified at Republic, '' Pentacentron sternhartae'', ''
Tetracentron hopkinsii ''Tetracentron hopkinsii'' is an extinct species of flowering plant in the family Trochodendraceae. The species is known from fossil leaves found in the early Eocene deposits of northern Washington state, United States and south Central Britis ...
'', and ''
Trochodendron nastae ''Trochodendron nastae'' is an extinct species of flowering plant in the family Trochodendraceae known from fossil leaves found in the early Eocene Ypresian stage Klondike Mountain Formation deposits of northern Washington state. ''T. nastae'' i ...
''. Additionally the species '' Trochodendron drachukii'' is known from related Kamloops group shales at 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 ...
near Cache Creek, British Columbia. Manchester ''et al.'' 2018B noted that ''Tr. drachukii'' is likely the fruits of ''Tr. nastae'', while ''Pe. sternhartae'' are likely the fruits of ''Te. hopkinsii''. If fossils of the fruits and foliage in attachment are found, that would bring the species count down to three whole plant taxa.


Description

The
fruiting bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
of ''Paraconcavistylon wehrii'' are simple
raceme A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...
s that taper from a wide base down to a tip under wide, with lengths of up to and over . Fruit capsules are born on
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 ...
s which curve upwards towards the raceme apex in a helical pattern and the longest preserved raceme has 33 attached fruits, though the specimen is missing both basal and apical sections. Given the length of the racemes, they likely hung down like a pendulum, with the fruits pointing downwards. Each of the smooth teardrop shaped fruits is between wide by tall. They have a slight thickening in the basal area where a ring of
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 ...
scars encircling the connection with the pedicel is, but are widest near the fruit apex. Between four and six long persistent styles are arranged just below the midpoint of the capsules and curving upwards towards the capsule apex. At the base of each style are swollen elliptical bulges that likely were
nectaries Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
. The fruits opened at the apex, with a four to six rayed star pattern formed by the dehiscence splits which run from just above the styles to the fruit apex where they join together. ''P. wehrii'' twigs have alternating leaves with terminal and axillary buds. The raceme is born from a
fork In cutlery or kitchenware, a fork (from la, furca 'pitchfork') is a utensil, now usually made of metal, whose long handle terminates in a head that branches into several narrow and often slightly curved tines with which one can spear foods ei ...
of the twig and leaf, as are the pointed buds. Typically the axillary buds are around long by while the terminal buds are larger at long by . Encircling the twig are between five and eight distinct terminal bud scars. The leaves have a narrow base connecting to the long petioles. They have regularly spaced teeth along the margin that are rounded and bearing glands, and a few specimens from the McAbee site also have distinct laminal lobes bracketing the petiole. The leaves have an overall range between long by , with an obovate outline. Like ''Trochodendron'' the leaves have a pinnate vein structure, with between eight and fifteen secondary veins that fork from the central main vein and arch towards the leaf apex before merging with the secondary above.


References

{{reflist, 30em Trochodendrales Plants described in 2020 Fossil taxa described in 2020 Ypresian plants of North America Extinct flora of North America Klondike Mountain Formation Prehistoric angiosperm genera Tranquille Formation