Macginitiea Gracilis SRIC SR 01-03-04 A Img1
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''Macginitiea'' is an extinct
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 ...
in the family
Platanaceae Platanaceae, the "plane-tree family", is a family of flowering plants in the order Proteales. The family consists of only a single extant genus ''Platanus'', with eight known species. The plants are tall trees, native to temperate and subtropic ...
ranging from the Late Paleocene to Late Eocene of North America, known from the
Clarno Formation Clarno may refer to: Locations * Clarno, Oregon * Clarno, Wisconsin ** Clarno (community), Wisconsin *Clarno Township, Lake County, South Dakota Lake County is a county in the U.S. state of South Dakota. As of the 2020 United States Census, t ...
of central
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
. The genus is strictly used to describe
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
, but has been found in close association with other fossil platanoid organs, which collectively have been used for whole plant reconstructions. ''Macginitiea'' and its associated organs are important as together they comprise one of the most well-documented and ubiquitous fossil plants, particularly in the Paleogene of North America.Myers, J. A. (2003). Terrestrial Eocene-Oligocene Vegetation and Climate in the Pacific Northwest. ''From Greenhouse to Icehouse: The Marine Eocene-Oligocene Transition. Columbia University Press, New York'', 171-185. Because paleobotanical material is often found in disarticulation, different species names are often used to refer to different organs (e.g. leaves, fruits, wood) even if those organs might have belonged to the same plant. When these organ species are considered together as a whole plant, the study is known as a whole plant reconstruction. Some localities have enough co-occurrences of different fossil plant organs that whole plant reconstructions are possible, one example being the ''
Cercidiphyllum ''Cercidiphyllum'' is a genus containing two species of plants, both commonly called katsura. They are the sole members of the monotypic family Cercidiphyllaceae. The genus is native to Japan and China and unrelated to ''Cercis'' (redbuds). Desc ...
''-like ''Joffrea'' from the Joffre Bridge locality of
Alberta, Canada Alberta ( ) is one of the thirteen provinces and territories of Canada, provinces and territories of Canada. It is part of Western Canada and is one of the three Canadian Prairies, prairie provinces. Alberta is bordered by British Columbia to t ...
. The earliest definite occurrences of ''Macginitiea'' have been observed from the Comstock Flora of southern
Willamette Valley The Willamette Valley ( ) is a long valley in Oregon, in the Pacific Northwest region of the United States. The Willamette River flows the entire length of the valley and is surrounded by mountains on three sides: the Cascade Range to the east, ...
, Oregon, which has been dated to be younger than 38-40 million years old (
Bartonian The Bartonian is, in the ICS's geologic time scale, a stage or age in the middle Eocene Epoch or Series. The Bartonian Age spans the time between . It is preceded by the Lutetian and is followed by the Priabonian Age. Stratigraphic definitio ...
), though similar ages are observed throughout the
Pacific Northwest The Pacific Northwest (sometimes Cascadia, or simply abbreviated as PNW) is a geographic region in western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though ...
, with occurrences around 38-39 Ma.


Description

''Macginitiea'' leaves are readily recognizable by their three to nine, equally spaced, palmate lobes. Like other Platanaceae, ''Macginitiea'' has palinactinodromous primary venation, a type of venation wherein primary veins fork multiple times. ''Macginitiea'' also has an inflated petiole base, which in modern counterparts encloses the underlying axillary buds for the next year, indicating their
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
nature. The branching of primary veins often occurs basally, rather than suprabasally (as encountered in some modern species). Although the secondary veins of modern ''
Platanus ''Platanus'' is a genus consisting of a small number of tree species native to the Northern Hemisphere. They are the sole living members of the family Platanaceae. All mature members of ''Platanus'' are tall, reaching in height. All except f ...
'' are usually straight, the secondary veins of ''Macginitiea'' are so regular and prominent compared to other platanoids that this recognizable "chevron" pattern has been considered a primary characteristic of the species.Wolfe, J. A., & Wehr, W. (1987). Middle Eocene dicotyledonous plants from Republic, northeastern Washington. Margins are usually entire in all ''Macginitiea'' species, but can sometimes have minor teeth. ''Macginitiea'' differs from modern ''
Platanus ''Platanus'' is a genus consisting of a small number of tree species native to the Northern Hemisphere. They are the sole living members of the family Platanaceae. All mature members of ''Platanus'' are tall, reaching in height. All except f ...
'' with its often greater number of lobes and narrower angle between adjacent primary veins. Different species of ''Macginitiea'' can be distinguished on the basis of lobe depth, extent of chevrons, prevalence of teeth, and partly lobe number. In total, 5 species of ''Macginitiea'' leaves have been described. Table adapted from Pigg & Stockey (1991).


Whole plant reconstructions


The Clarno Plane

The "Clarno Plane" was established as an informal name to refer to the whole plant recognized from five fossil species: ''Macginitiea angustiloba'' (
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
), '' Plataninium haydenii'' (
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin th ...
), '' Macginicarpa glabra'' ( infructescences), '' Platananthus synandrus'' ( staminate inflorescences), and '' Macginistemon mikanoides'' (isolated
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 ...
clusters). The Clarno Plane is known from the west coast of
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
across several states, including central
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
,
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
, and northern
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 ...
. Like the modern ''Platanus,'' ''Macginicarpa'' has clusters of five
carpels Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
per
floret This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...
. However, in striking contrast to the modern, ''Macginicarpa'' flowers have a well-developed
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 ...
and fruits lack the prominent dispersal hairs characteristic of the modern ''Platanus''. Modern carpel numbers are more variable than the consistent five of ''Macginicarpa,'' ranging from four to nine. Staminate flowers (''Platananthus synandrus'') have an even more developed perianth than pistillate flowers, but similarly have more consistent numbers of parts than the modern, with 5
stamens 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 ...
per
floret This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...
. ''P. synandrus''
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
appears to be smaller than pollen from modern ''Platanus''. ''Platananthus synandrus'' is also distinctive from extant ''Platanus'' for the elongation of its
connectives In logic, a logical connective (also called a logical operator, sentential connective, or sentential operator) is a logical constant. They can be used to connect logical formulas. For instance in the syntax of propositional logic, the binary co ...
, extensions of filament tissue that cover or divide an
anther 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 filam ...
. In ''Platananthus'' and in modern ''Platanus'',
peltate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
(shield-like) connectives cover the tops of anthers, but the connectives of ''Platananthus'' are 4 to 5 times the length of the modern. Stamens are connate (fused) within each floret, causing them to be shed in clusters of stamen bundles, rather than one at a time as in modern ''Platanus'' species. Stamen bundles associated with ''Macginitiea'' have been put under the genus ''Macginistemon''. ''Plataninium'' wood is similar, but has wider rays and more scalariform (ladder-like)
perforations A perforation is a small hole in a thin material or web. There is usually more than one perforation in an organized fashion, where all of the holes collectively are called a ''perforation''. The process of creating perforations is called perfor ...
than that of recent species. As of 1986, Dr. Steven Manchester said that "The Clarno Plane is currently the most completely documented fossil angiosperm species, known morphologically and anatomically from wood, leaves, pistillate and staminate inflorescences, fruits, and pollen." Associations of multiple organs of the Clarno Plane in various combinations have been found in "more than ten localities" throughout western North America, as of 2008. Other platanoid leaves have since been found in association with reproductive structures, including leaves such as '' Platimeliphyllum'', '' Ettingshausenia'', '' Evaphyllum'', '' Platanus neptuni'', '' Platanus nobilis'', and ''
Sapindopsis ''Sapindopsis'' ("''Sapindus''-like") is an extinct form genus for leaves of the Cretaceous Period, originally considered similar to soapberry. Associated reproductive structures now suggest it was more closely related to planes and sycamores of ...
''.


The Joffre Plane

The Paleocene fossil leaf species ''Platanus nobilis'' was established as a species intermediate between ''Macginitiea'' and modern ''Platanus''. However, differences between ''P. nobilis'' and ''Macginitiea'' were later considered too minor to justify placing ''P. nobilis'' in a different genus, particularly since ''P. nobilis'' was associated with ''Macginicarpa'' inflorescences. As such, ''P. nobilis'' was reassigned to ''Macginitiea nobilis.'' The "Joffre Plane" as a whole plant reconstruction includes leaves from ''Macginitiea nobilis'', pistillate inflorescences and infructescences from ''Macginicarpa manchesteri'', and staminate inflorescences of ''Platananthus speirsae''. ''Macginitiea nobilis'' is set apart from other ''Macginitiea'' species by its fewer number of lobes (usually 3, instead of 5-9) and less distinct "chevron" venation pattern. ''M. nobilis'' has been found in various stages of development from the Joffre Bridge locality, from
seedlings A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embry ...
with cotyledonous leaves to mature, true, trilobate leaves.


Discovery and naming

The genus ''Macginitiea'' was named after the prominent paleobotanist Harry Dunlap MacGinitie, who was the first person to suggest the close relationship of ''Macginitiea'' to the family
Platanaceae Platanaceae, the "plane-tree family", is a family of flowering plants in the order Proteales. The family consists of only a single extant genus ''Platanus'', with eight known species. The plants are tall trees, native to temperate and subtropic ...
. Though fossils of ''Macginitiea'' were originally classified under the genus ''
Aralia ''Aralia'' , or spikenard, is a genus of the family Araliaceae, consisting of 68 accepted species of deciduous or evergreen trees, shrubs, and rhizomatous herbaceous perennials. The genus is native to Asia and the Americas, with most species occu ...
,'' MacGinitie noticed that the venation patterns of ''Aralia angustiloba'' more closely resembled that of ''
Platanus ''Platanus'' is a genus consisting of a small number of tree species native to the Northern Hemisphere. They are the sole living members of the family Platanaceae. All mature members of ''Platanus'' are tall, reaching in height. All except f ...
'' rather than ''Aralia'', and reclassified ''Aralia angustiloba'' as ''Platanophyllum angustiloba.'' However, Wolfe & Wehr found that the genus ''Platanophyllum'' was problematic, and therefore made the new genus ''Macginitiea'' for these leaves.


Ecology

''Macginitiea'' has been known to be the dominant element in many
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' ...
floras. The Clarno Plane is thought to have occupied open, riparian habitats, similar to that of modern ''Platanus''. They also have been found several times in co-occurrence with ''
Cercidiphyllum ''Cercidiphyllum'' is a genus containing two species of plants, both commonly called katsura. They are the sole members of the monotypic family Cercidiphyllaceae. The genus is native to Japan and China and unrelated to ''Cercis'' (redbuds). Desc ...
''-like plants. The association with ''Cercidiphyllum'' suggests that the Clarno Plane was tolerant of full sunlight and poorly developed soils.


References


External links

* {{Taxonbar, from=Q21396968 Platanaceae Eocene plants of North America Eocene United States Paleogene geology of Oregon Eocene genus first appearances Eocene extinctions Clarno Formation