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''Florissantia'' is an extinct genus 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 ...
s in the
Malvaceae Malvaceae, or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include okra, cotton, cacao and durian. There are also some genera containing familiar ...
subfamily Sterculioideae known from western North America and far eastern
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 are ...
. Flower,
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
, and pollen
compression fossil A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way. The reason f ...
s have been found in formations ranging between the Early Eocene through to the
Early Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/ Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...
periods. The type species is ''Florissantia speirii'' and three additional species are known, ''Florissantia ashwillii'', ''Florissantia quilchenensis'', and ''Florissantia sikhote-alinensis''.


Distribution

Fossils of ''Florissantia'' have been found in Early Eocene to Middle Oligocene localities, including fossil beds in
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S. ...
,
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, ...
,
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
,
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columb ...
,
Oregon Oregon () is a state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington, while the Snake River delineates much of its eastern boundary with Idaho. T ...
,
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
,
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 o ...
,
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the s ...
and the
Russian far east The Russian Far East (russian: Дальний Восток России, r=Dal'niy Vostok Rossii, p=ˈdalʲnʲɪj vɐˈstok rɐˈsʲiɪ) is a region in Northeast Asia. It is the easternmost part of Russia and the Asian continent; and is admin ...
. The type species ''F. speirii'' is known from a number of formations in the west, with the type locality being the
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones an ...
of Colorado. The formation is composed of successive lake deposits resulting from a volcanic debris flow damming a valley. When the species was described, the Florissant Formation was considered to be
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
in age, based on the flora and fauna preserved. Successive research and fossil descriptions moved the age older and by 1985 the formation had been reassigned to an Oligocene age. Further refinement of the formation's age using radiometric dating of sanidine crystals has resulted in an age of placing the formation in the Priabonian stage of the Late Eocene. A tentative report of the species from the Katalla Formation,
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S. ...
was made by Jack Wolfe (1977) based on a single specimen. ''F. quilchenensis'' has been recovered from four locations in the Okanagan highlands, the Falklands and McAbee sites near
Cache Creek, British Columbia Cache Creek is a historic transportation junction and incorporated village northeast of Vancouver in British Columbia, Canada. It is on the Trans-Canada Highway in the province of British Columbia at a junction with Highway 97. The same in ...
, the Coldwater Beds Quilchena site near Quilchena, British Columbia and the Klondike Mountain Formation in Republic, Washington, northern
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. T ...
. The Okanagan highlands are aged between for the Quilchena site to for the Klondike Mountain Formations Tom Thumb Tuff member. The youngest ''F. quilchenensis'' occurrence is from the
early Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/ Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...
Gumboot Mountain Flora of southwestern Washington. Additionally ''Florissantia'' sp. fossils have also been reported from Okanagan highlands sites in the
Allenby Formation The Allenby formation is a sedimentary rock formation in British Columbia which was deposited during the Ypresian stage of the Early Eocene. It consists of conglomerates, sandstones with interbedded shales and coal. The shales contain an abunda ...
near Princeton, the Horsefly Shales near Horsefly, and the Driftwood Shales near Smithers, British Columbia, without indication of specie affinity. ''F. ashwillii'' is confined to the Middle to Late Eocene Clarno Formation, Latest Eocene Goshen flora and Early Oligocene
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 ...
of central Oregon. The type locality was designated by Manchester (1992) as the Sheep Rock Creek locality, which represents an isolated pond deposit likely correlating to Clarno Formation based on a palm frond fossil found at the site. The species also is present in the western Oregon "Goshen Flora", which is part of the Fisher Formation The youngest occurrences of ''F. ashwillii'' are confined to the Summer Spring shales locality around Grey Butte and thought to correlate with Early Oligocene strata of the John Day Formation. The Amgu flora, the type locality for ''F. sikhote-alinensis'', represents outcrops of the Khutsin Formation exposed in the
Sikhote Alin The Sikhote-Alin (russian: Сихотэ́-Али́нь, , , ) is a mountain range in Primorsky and Khabarovsk Krais, Russia, extending about to the northeast of the Russian Pacific seaport of Vladivostok. The highest summits are Tordoki Yani a ...
mountains on the Russian far-east coast. The flora was reported by Manchester (1999) as Miocene, however more detailed work on the stratigraphy of the region moved the age to Early Oligocene as noted by Kvaček ''et al'' (2005), Peng ''et al'' (2011) reported a slightly older late Eocene or Early Oligocene age which was followed by Archibald and Rasnitsyn (2018) who list the site as likely Priabonian or perhaps
Rupelian The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/ Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chattian ...
.


History & classification

Fossils now placed in ''Florissantia'' were first reported by
Leo Lesquereux Charles Léo Lesquereux (November 18, 1806 – October 25, 1889) was a Swiss-born bryologist and a pioneer of American paleobotany who studied the formation of peat bogs. Career Lesquereux was born in the town of Fleurier, located in the canto ...
( 1883) from the
Florissant Formation The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones an ...
and described as the morning glory family species '' Porana speirii''. Additional fossils were described by Knowlton (1916) as another species, ''Porana similis'' as well as a third species, which he placed into the new genus ''Florissantia'' as ''Florissantia physalis''. Calyxes of ''F. quilchenensis'' were first reported by Berry (1929), who identified them as the Miocene ''
Hydrangea ''Hydrangea'', () commonly named the hortensia, is a genus of over 75 species of flowering plants native to Asia and the Americas. By far the greatest species diversity is in eastern Asia, notably China, Korea, and Japan. Most are shrubs tall, ...
'' species '' Hydrangea bendirei''. The Republic fossils were later included by
Brown Brown is a color. It can be considered a composite color, but it is mainly a darker shade of orange. In the CMYK color model used in printing or painting, brown is usually made by combining the colors orange and black. In the RGB color model us ...
(1935) in ''Porana speiri''. Mathewes and Brooke (1971) described a solitary flower as the new species ''Holmskioldia quilchenensis'' from a fossil found at the Quilchena site in British Columbia, which Manchester (1992) redescribed based on a larger selection of specimens from Republic, and moved the species into ''Florissantia''. The first instance of ''F. ashwillii'' was published by
Chaney Chaney is a surname. Notable people with the surname include: * Cassius Chaney (born 1987), American boxer * Charles "Bubba" Chaney (born 1946), American politician * Chris Chaney, American musician * Darrel Chaney, American baseball player * Do ...
and Sanborn (1933) who included the flowers as in their species '' Viburnum palmatum''. Manchester (1992) described the ''Florissantia'' species based on a series of 45 fossils found in north central Oregon and noted that the 1933 flower was not related to the leaf holotype fossil of ''V. palmatum''. The ''F. ashwillii'' holotype specimen, UF 11740, was a part of the University of Florida collections at the time of description, with additional fossils examined from the
University of California Museum of Paleontology The University of California Museum of Paleontology (UCMP) is a paleontology museum located on the campus of the University of California, Berkeley. The museum is within the Valley Life Sciences Building (VLSB), designed by George W. Kelham and ...
collections. Manchester chose the specific epithet ''ashwillii'' as a patronym honoring Melvin S. Ashwill who collected many of the specimens examined.


Description

All ''Florissantia'' species share a distinct set of physical traits in common. The flowers are
radially symmetrical Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
shallowly bell shaped, with large
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coine ...
s fused up to 50% of their length and born on long
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. The sepals display a distinct reticulate venation radiating from the center of each sepal with the crossveins forming rectangular to polygonal meshes of smaller veins. Between five and seven major veins run from the sepal base apically before loping towards the apex and fusing together. Additional large veins run to the point of each sinus, and supporting several secondary veins. The basal calyx is thicker than the sepal tissue and often hairy with the small thick petals forming a corolla around the calyx base. The flowers have an expanded androgynophore from which the ovary arises. A single
style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
extends from the center of the ovary though the surrounding
androecium 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 fila ...
to above the corolla of five stamens. The stamen filaments fork once near the tip, and host ten total anthers with elongated to globose outlines. The pollen grains are slightly flattened spheres, with three to four short colpi apertures on the outer surface


''Florissantia ashwillii''

''Florissantia ashwillii'' calyxes are moderate in size with a diameter of , those of ''F. quichenensis'' are known to be smaller, while those of ''F. speirii'' range notably larger. ''F. ashwilli'' is noted for having deeply indented lobe sinuses which extend between 35% to 60% towards the center of the corolla from the margins. The pedicels are at least long, with a hairy basal area of the corolla, and pollen smaller than that of ''F. speirii''.


''Florissantia quilchenensis''

''F. quilchenensis'' is calyxes range between , with some of the smallest recorded specimens in ''Florissantia''. unlike the other three species, the calyx lobes range from approximately 33% indentation to being fully absent, resulting in a circular calyx. At the time of description, only three specimens had been recovered in which the stamens were present and approximately ten anthers or half anthers are preserved on the most complete of the specimens. The
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 are longer than those in ''F. ashwillii'', being between long and thick. The petals are around and are arranged in alternation with the Calyx lobes, when the lobes are present. The style is long and extends above the stamens. Rather than having globose anthers, such as seen in ''F. speirii'', the long anthers are elongated like those seen in ''F. ashwillii''.


''Florissantia sikhote-alinensis''

''Florissantia sikhote-alinensis'' was first described as ''Porana '' by Kryshtofov (1921). The fossil pictured in plate 2 fig. 7 of the paper was subsequently lost and was unavailable for reexamination by Manchester in 1998.


''Florissantia speirii''

''F. speirii'' calyxes range between having some of the largest calyxes in ''Florissantia''. The calyxes show poorly developed to well developed lobes that indent between 20%–35% from the outer margins and in flowers with poorly developed indentation, the calyx outline is pentagonal instead of rounded. The androgynophore has a rounded pentagonal to circular cross-section and an approximately long style. A tepal corolla has been found attached to a single fossil from the
Florissant Fossil Beds National Monument The Florissant Formation is a sedimentary geologic formation outcropping around Florissant, Teller County, Colorado. The formation is noted for the abundant and exceptionally preserved insect and plant fossils that are found in the mudstones and ...
collections.


Paleoecology

The structure of ''Florissantia'' flowers suggests the genus was
pollinated Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
by insects or birds. With the long slender pedicels, the flowers likely hung downwards from the plant in an
pendant A pendant is a loose-hanging piece of jewellery, generally attached by a small loop to a necklace, which may be known as a "pendant necklace". A pendant earring is an earring with a piece hanging down. Its name stems from the Latin word ' ...
like manner, with dense hairs along the flower bases acting as
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 ...
. These are features seen in a number of modern Malvaceae genera. Additionally the stamens and styles are thicker and robust, features often associated with bird and bat pollinators, indicating ''Florissantia'' may have been a transitional genus between insect pollination and bat or bird pollination. It is possible the caylxes did not grow to full size until after pollination, with the young flowers being smaller and less showy. It is likely the fruits matured into a samara with the calyx acting as a wing around the fruit.


References


External links

* {{Taxonbar, from=Q5461909 Sterculioideae Eocene plants Oligocene plants Eocene life of North America Oligocene life of North America Prehistoric angiosperm genera Malvaceae genera Prehistoric plants of North America Allenby Formation Coldwater Beds Florissant Formation Horsefly Shales Klondike Mountain Formation Tranquille Formation Clarno Formation