Calamopityaceae Temporal range: | |
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Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | † Pteridospermatophyta |
Order: | † Calamopityales Němejc, 1963 |
Family: | † Calamopityaceae Solms, 1896 |
Synonyms | |
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Calamopityaceae is the largest family of the division of extinct seed-bearing plants (spermatophytes) known as Pteridospermatophyta. [1] [2] It is the only family in the monotypic order Calamopityales. [3] This family is characterized by its petioles and specific wood pattern, and it grew only in the Paleozoic era, specifically in North America and Europe. [1] [4] [5] Three form genera within the family are diagnosed by their stem structure: Calamopitys , Stenomyelon , and Diichinia . [1] It was named by Solms-Laubach in 1896. Since then, its genera have been added to and grouped differently. [1]
Calamopityaceae is the largest family in Pteriodspermatophyta. [2] This family is composed of gymnosperms, and because of their stem structure discovered through fossil rocks, they are considered to be in this division. [6] However, nothing is known of their reproductive organs, but they are classified as seed plants based on their similarities to the Lyginopteridaceae and Medullasaceae families within Pteriodspermatophyta. [4] Calamopityaceae resemble Lyginopteridaceae and Medullasaceae in the monoxylic wood structures in their stem; this structure suggests the stem (diameter less than 1.5 cm) was narrow during the Calamopityaceae plant lifetime. [5] [6] Only some petiole tissue has been found; it is classified to be of the genus Kalymma and suggests the plant had large fronds. [4] [5] To identify a genus within this family, this petiole structure and monoxylic wood must be present, as well as a much larger cortex than vascular cylinder. [4] No fossil evidence has been found to describe on their seed and pollen (reproductive) organs, and therefore the species within this family show more variance than other families. [1] [5]
Gymnosperms, including those that are extinct, can be classified by their wood: monoxylic vs pycnoxylic. [7] Monoxylic wood is soft and spongy and has a large pith and cortex. [6] Pycnoxylic wood, which is more dense with less pith and cortex, is more commercially used. [7] The three genera of Calamopityaceae, Calamopitys, Stenomyelon, and Diichnia, show monoxylic wood stem patterns, and this is considered to be an essential classification of the family Calamopityaceae (hence why Bilignea, Eristophyton, Endoxylon, and Shenoxylon were removed from this family). [1]
Calamopityaceae fossils have been found in North America and Europe, and they have been dated back to the Paleozoic era, specifically the Upper Devonian and Lower Carboniferous (Mississippian) periods. [5] [6] Being from this early period, Calamopityaceae are significant as an example of some of the earliest seed plants and ancestors of angiosperms. [4] [5]
In 1856, the Austrian paleontologist, Franz Joseph Andreas Nicolaus Unger, was the first to find Calamopitys, a genus of Calamopityaceae. [1] This genus, which later was the root for the family name, was found in the Thuringian Forest. [1] Forty years later, the family was named Calamopityaceae by Solms-Laubach.
Though the original genera, Calamopitys, Stenomyelon, and Diichnia, still remain under this family classification, there have been historical additions to these groupings. Because the family is defined loosely on stem structure with nothing known about the foliage and reproductive structure, different genera have been added and removed from this family. [5] Four more genera, Bilignea , Eristophyton , Endoxylon , and Sphenoxylon, were added to the family in 1936. These genera were classified by their pycnoxylic secondary wood pattern. In 1953, they were removed from the family with the intention of keeping the family composed of genera with monoxylic secondary wood. [1]
Like many "seed ferns", the affinities of the Calamopityales are unclear. Meyen (1984) placed them within the class Ginkgoopsida, which also includes ginkgos. Apart from Calamopitys and Stenomyelon, he also placed the genera Lyrasperma , Deltasperma , Eosperma , and Kalymma within Calamopityaceae. [8]
A classification scheme by Doweld (2001) considered Calamopityales to be an incertae sedis order within the phylum "Moresnetiophyta". Doweld also listed Buteoxylaceae Barnard & Long 1973 [9] as a synonym of Calamopityaceae, implying that its constituent fossil wood genera ( Buteoxylon and Triradioxylon ) should be referred to Calamopityaceae. This would additionally require treating the order Buteoxylonales as a synonym of Calamopityales. [10] Stenomyelon has been compared favorably to Buteoxylon in more recent studies as well. [11]
Three genera of fossil wood are currently classified as belonging to the family Calamopityaceae, and their differences are distinguished by their decreasing primary xylem from Stenomyelon, to Calamopitys, to Diichnia. [12] Additionally, Galtiera , Triichnia , and Bostonia are classified within the family by some authors, [13] [14] and Faironia [15] and Chapelia [16] may also have affinities with the family.
Type species - Calamopitys saturnii [1]
There are six species within this genus, and it has the most species of any Calamopityaceae genera. [6] Although Eristophyton is sometimes considered to be a subgenus under this genus, the distinction between pycnoxylic and monoxylic secondary wood maintains these genera as separate. [6] In terms of structure, these plants under this genera have narrow stems with diameter 2–3 cm (or larger in C. embergeri and C. schweitzeri). [5]
Type species - Stenomyelon tuedianum [1]
Originally known as "Tweed Mill fossil", this genus consists of fewer species than Calamopitys, but in addition to S. tuedianum, species include S. primaevum, S. heterangioides, and S. muratum. [1]
Type species - Diichnia kentuckiensis [1]
Species of this genus are classified based on the characteristics of the smallest primary xylem of Calamopityaceae and a five-angled pith, as seen in the stem cross-section. [6] These characteristics separate this genus from the other genera.
Non-wood form genera referred to Calamopityaceae include: [17]
Araucariaceae – also known as Araucarians – is a family of coniferous trees, with three living genera, Araucaria, Agathis, and Wollemia. While the family was a common component of the flora globally during the Jurassic, Cretaceous and Paleogene periods, in their native distribution they are now largely confined to the Southern Hemisphere, except for a few species of Agathis in Malesia.
Paleobotany, also spelled as palaeobotany, is the branch of botany dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeography), and the evolutionary history of plants, with a bearing upon the evolution of life in general. A synonym is paleophytology. It is a component of paleontology and paleobiology. The prefix palaeo- or paleo- means "ancient, old", and is derived from the Greek adjective παλαιός, palaios. Paleobotany includes the study of terrestrial plant fossils, as well as the study of prehistoric marine photoautotrophs, such as photosynthetic algae, seaweeds or kelp. A closely related field is palynology, which is the study of fossilized and extant spores and pollen.
Glossopteris is the largest and best-known genus of the extinct Permian order of seed plants known as Glossopteridales. The name Glossopteris refers only to leaves, within the framework of form genera used in paleobotany.
Pteridospermatophyta, also called "pteridosperms" or "seed ferns" are a polyphyletic grouping of extinct seed-producing plants. The earliest fossil evidence for plants of this type are the lyginopterids of late Devonian age. They flourished particularly during the Carboniferous and Permian periods. Pteridosperms declined during the Mesozoic Era and had mostly disappeared by the end of the Cretaceous Period, though Komlopteris seem to have survived into Eocene times, based on fossil finds in Tasmania.
Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales were amongst the most common seed plants of the Mesozoic, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.
Torreya is a genus of conifers comprising six or seven species placed in the family Taxaceae, though sometimes formerly placed in Cephalotaxaceae. Four species are native to eastern Asia; the other two are native to North America. They are small to medium-sized evergreen trees reaching 5–20 m, rarely 25 m, tall. Common names include nutmeg yew.
Cheirolepidiaceae is an extinct family of conifers. They first appeared in the Triassic, and were a diverse and common group of conifers during most of the Mesozoic era, primarily at low latitudes, where they often formed a dominant element of the vegetation. They are united by the possession of a distinctive pollen type assigned to the form genus Classopollis The name Frenelopsidaceae or "frenelopsids" has been used for a group of Cheirolepidiaceae with jointed stems, thick internode cuticles, sheathing leaf bases and reduced free leaf tips. The leaf morphology has been noted as being similar to that of halophyte Salicornia. Several members of the family appear to have been adapted for semi-arid and coastal settings, with a high tolerance of saline conditions. Cheirolepidiaceae disappeared from most regions of the world during the Cenomanian-Turonian stages of the Late Cretaceous, but reappeared in South America during the Maastrichtian, the final stage of the Cretaceous, increasing in abundance after the K-Pg extinction and being a prominent part of the regional flora during the Paleocene, before going extinct. Survival into the Paleocene in North America and China has also been suggested based on pollen.
Psaronius is an extinct genus marattialean tree fern which grew to 10m in height, and is associated with leaves of the organ genus Pecopteris and other extinct tree ferns. Originally, Psaronius was a name for the petrified stems, but today the genus is used for the entire tree fern. Psaronius tree fern fossils are found from the Carboniferous through the Permian.
The Medullosales is an extinct order of pteridospermous seed plants characterised by large ovules with circular cross-section and a vascularised nucellus, complex pollen-organs, stems and rachides with a dissected stele, and frond-like leaves. Their nearest still-living relatives are the cycads.
The Lyginopteridales are an extinct group of seed plants known from the Paleozoic. They were the first plant fossils to be described as pteridosperms and, thus, the group on which the concept of pteridosperms was first developed; they are the stratigraphically oldest-known pteridosperms, occurring first in late Devonian strata; and they have the most primitive features, most notably in the structure of their ovules. They probably evolved from a group of Late Devonian progymnosperms known as the Aneurophytales, which had large, compound frond-like leaves. The Lyginopteridales became the most abundant group of pteridosperms during Mississippian times, and included both trees and smaller plants. During early and most of middle Pennsylvanian times the Medullosales took over as the more important of the larger pteridosperms but the Lyginopteridales continued to flourish as climbing (lianescent) and scrambling plants. However, later in Middle Pennsylvanian times the Lyginopteridales went into serious decline, probably being out-competed by the Callistophytales that occupied similar ecological niches but had more sophisticated reproductive strategies. A few species continued into Late Pennsylvanian times, and in Cathaysia and east equatorial Gondwana they persisted into the Late Permian, but subsequently became extinct. Most evidence of the Lyginopteridales suggests that they grew in tropical latitudes of the time, in North America, Europe and China.
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Dicroidium is an extinct genus of fork-leaved seed plants. It is the archetypal genus of the corystosperms, an extinct group of seed plants, often called "seed ferns", assigned to the order Corystospermales or Umkomasiales. Species of Dicroidium were widely distributed and dominant over Gondwana during the Triassic. Their fossils are known from South Africa, the Arabian Peninsula, Australia, New Zealand, South America, Madagascar, the Indian subcontinent and Antarctica.
The Callistophytaceae was a family of seed ferns (pteridosperms) from the Carboniferous and Permian periods. They first appeared in late Middle Pennsylvanian (Moscovian) times, 306.5–311.7 million years ago (Ma) in the tropical coal forests of Euramerica, and became an important component of Late Pennsylvanian vegetation of clastic soils and some peat soils. The best known callistophyte was documented from Late Pennsylvanian coal ball petrifactions in North America.
The Peltaspermales are an extinct order of seed plants, often considered "seed ferns". They span from the Late Carboniferous to the Early Jurassic or the Jurassic-Cretaceous Boundary. It includes at least one valid family, Peltaspermaceae, which spans from the Permian to Early Jurassic, which is typified by a group of plants with Lepidopteris leaves, Antevsia pollen-organs, and Peltaspermum ovulate organs, though the family now also includes other genera like Peltaspermopsis, Meyenopteris and Scytophyllum. Along with these, two informal groups of uncertain taxonomic affinities exist, each centered around a specific genus ; Supaia and Comia, known from the Early Permian of the Northern Hemisphere, especially of North America. Both the "Comioids" and the "Supaioids" are associated with the peltaspermacean ovulate organ Autunia. The Late Triassic-Middle Jurassic genus Pachydermophyllum may also have affinities to the peltasperms.
Corystosperms are a group of extinct seed plants belonging to the family Corystospermaceae assigned to the order Corystospermales or Umkomasiales. They were first described based on fossils collected by Hamshaw Thomas from the Burnera Waterfall locality near the Umkomaas River of South Africa. Corystosperms are typified by a group of plants that bore forked Dicroidium leaves, Umkomasia cupulate ovulate structures and Pteruchus pollen organs, which grew as trees that were widespread over Gondwana during the Middle and Late Triassic. Other fossil Mesozoic seed plants with similar leaf and/or reproductive structures have also sometimes been included within the "corystosperm" concept sensu lato, such as the "doyleoids" from the Early Cretaceous of North America and Asia. A potential corystosperm sensu lato, the leaf genus Komlopteris, is known from the Eocene of Tasmania, around 53-50 million years old, over 10 million years after the Cretaceous–Paleogene extinction event.
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