Pteridospermatophyta Late | |
---|---|
Fossil seed fern leaves of Neuropteris (Medullosales) from the Late Carboniferous of northeastern Ohio. | |
Life restoration of Lepidopteris (Peltaspermales) | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Spermatophytes |
Division: | † Pteridospermatophyta |
Groups included | |
| |
Excluded | |
| |
Synonyms | |
Pteridospermatopsida |
Pteridospermatophyta, also called "pteridosperms" or "seed ferns" are a polyphyletic [1] grouping of extinct seed-producing plants. The earliest fossil evidence for plants of this type are the lyginopterids of late Devonian age. [2] 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. [3]
With regard to the enduring utility of this division, many palaeobotanists still use the pteridosperm grouping in an informal sense to refer to the seed plants that are not angiosperms, coniferoids (conifers or cordaites), ginkgophytes or cycadophytes (cycads or bennettites). This is particularly useful for extinct seed plant groups whose systematic relationships remain speculative, as they can be classified as pteridosperms with no valid implications being made as to their systematic affinities. Also, from a purely curatorial perspective the term pteridosperms is a useful shorthand for describing the fern-like fronds that were probably produced by seed plants, which are commonly found in many Palaeozoic and Mesozoic fossil floras.
The concept of pteridosperms goes back to the late 19th century when palaeobotanists came to realise that many Carboniferous fossils resembling fern fronds had anatomical features more reminiscent of the modern-day seed plants, the cycads. In 1899 the German palaeobotanist Henry Potonié coined the term "Cycadofilices" ("cycad-ferns") for such fossils, suggesting that they were a group of non-seed plants intermediate between the ferns and cycads. [4] Shortly afterwards, the British palaeobotanists Frank Oliver and Dukinfield Henry Scott (with the assistance of Oliver's student at the time, Marie Stopes) made the critical discovery that some of these fronds (genus Lyginopteris ) were associated with seeds (genus Lagenostoma ) that had identical and very distinctive glandular hairs, and concluded that both fronds and seeds belonged to the same plant. [5] Soon, additional evidence came to light suggesting that seeds were also attached to the Carboniferous fern-like fronds Dicksonites , [6] Neuropteris [7] and Aneimites. [8] Initially it was still thought that they were "transitional fossils" intermediate between the ferns and cycads, and especially in the English-speaking world they were referred to as "seed ferns" or "pteridosperms". Today, despite being regarded by most palaeobotanists as only distantly related to ferns, these spurious names have nonetheless established themselves. Nowadays, four orders of Palaeozoic seed plants tend to be referred to as pteridosperms: Lyginopteridales, Medullosales, Callistophytales and Peltaspermales, with "Mesozoic seed ferns" including the Petriellales, Corystospermales and Caytoniales. [9]
Their discovery attracted considerable attention at the time, as the pteridosperms were the first extinct group of vascular plants to be identified solely from the fossil record. In the 19th century the Carboniferous Period was often referred to as the "Age of Ferns" but these discoveries during the first decade of the 20th century made it clear that the "Age of Pteridosperms" was perhaps a better description.
During the 20th century the concept of pteridosperms was expanded to include various Mesozoic groups of seed plants with fern-like fronds, such as the Corystospermaceae. Some palaeobotanists also included seed plant groups with entire leaves such as the Glossopteridales and Gigantopteridales, which was stretching the concept. In the context of modern phylogenetic models, [10] the groups often referred to as pteridosperms appear to be liberally spread across a range of clades, and many palaeobotanists today would regard pteridosperms as little more than a paraphyletic 'grade-group' with no common lineage.[ clarification needed ] One of the few characters that may unify the group is that the ovules were borne in a cupule, a group of enclosing branches, but this has not been confirmed for all "pteridosperm" groups.
It has been speculated that some seed fern groups may be close to the ancestry of flowering plants (angiosperms). A 2009 study concluded that "phylogenetic analysis techniques have surpassed the hard data needed to formulate meaningful phylogenetic hypotheses" regarding the relationships of "seed ferns" to living plant groups. [11]
Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or female. Cycads vary in size from having trunks only a few centimeters to several meters tall. They typically grow slowly and have long lifespans. Because of their superficial resemblance to palms or ferns, they are sometimes mistaken for them, but they are not closely related to either group. Cycads are gymnosperms (naked-seeded), meaning their unfertilized seeds are open to the air to be directly fertilized by pollination, as contrasted with angiosperms, which have enclosed seeds with more complex fertilization arrangements. Cycads have very specialized pollinators, usually a specific species of beetle. Both male and female cycads bear cones (strobili), somewhat similar to conifer cones.
The gymnosperms are a group of seed-producing plants that include conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, and literally means 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, and Ginkgo. The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.
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.
Archaeopteris is an extinct genus of progymnosperm tree with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous, the oldest fossils being 385 million years old, and had global distribution.
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.
Coal forests were the vast swathes of swamps and riparian forests that covered much of the land on Earth's tropical regions during the late Carboniferous (Pennsylvanian) and Permian periods. As plant matter from these forests decayed, enormous deposits of peat accumulated, which later became buried and converted into coal over the subsequent eras.
Macrozamia concinna is a part of the plant family, Zamiaceae. It originates from a division of Cycadophyta which encompasses the complete species of cycads. M. concinna is primarily habituated in New South Wales, Australia and maintains a distinct appearance allowing it to be easily identifiable from other cycads. M. concinna also implements a unique method of reproduction to fertilise its offsprings, as opposed to the common method of wind pollination. This difference in reproduction mechanisms has survived throughout the ages of prehistoric cycad species and M. concinna continues to procreate with it.
The euphyllophytes are a clade of plants within the tracheophytes. The group may be treated as an unranked clade, a division under the name Euphyllophyta or a subdivision under the name Euphyllophytina. The euphyllophytes are characterized by the possession of true leaves ("megaphylls"), and comprise one of two major lineages of extant vascular plants. As shown in the cladogram below, the euphyllophytes have a sister relationship to the lycopodiophytes or lycopsids. Unlike the lycopodiophytes, which consist of relatively few presently living or extant taxa, the euphyllophytes comprise the vast majority of vascular plant lineages that have evolved since both groups shared a common ancestor more than 400 million years ago. The euphyllophytes consist of two lineages, the spermatophytes or seed plants such as flowering plants (angiosperms) and gymnosperms, and the Polypodiophytes or ferns, as well as a number of extinct fossil groups.
This article attempts to place key plant innovations in a geological context. It concerns itself only with novel adaptations and events that had a major ecological significance, not those that are of solely anthropological interest. The timeline displays a graphical representation of the adaptations; the text attempts to explain the nature and robustness of the evidence.
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.
A seed plant or spermatophyte, also known as a phanerogam or a phaenogam, is any plant that produces seeds. It is a category of embryophyte that includes most of the familiar land plants, including the flowering plants and the gymnosperms, but not ferns, mosses, or algae.
Callistophytales is an extinct order of spermatophytes which lived from the Pennsylvanian to Permian periods. They were mainly scrambling and lianescent (vine-like) plants found in the wetland "coal swamps" of Euramerica and Cathaysia. Like many other early spermatophytes, they could be described as "seed ferns", combining ovule-based reproduction with pinnate leaves superficially similar to modern ferns.
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.
Emplectopteridaceae is an extinct family of pteridosperms known mainly from Permian floras of the Cathaysian Realm. They were mostly shrubby plants with a scrambling or upright habit, and favoured a range of habitats from arid to moist or even aquatic.
Asterotheca is a genus of seedless, spore-bearing, vascularized ferns dating from the Carboniferous of the Paleozoic to the Triassic of the Mesozoic.
Macroneuropteris is a genus of Carboniferous seed plants in the order Medullosales. The genus is best known for the species Macroneuropteris scheuchzeri, a medium-size tree that was common throughout the late Carboniferous Euramerica. Three similar species, M. macrophylla, M. britannica and M. subauriculata are also included in the genus.
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.
Calamopityaceae is the largest family of the division of extinct seed-bearing plants (spermatophytes) known as Pteridospermatophyta. It is the only family in the monotypic order Calamopityales. 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. Three form genera within the family are diagnosed by their stem structure: Calamopitys, Stenomyelon, and Diichinia. It was named by Solms-Laubach in 1896. Since then, its genera have been added to and grouped differently.
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.
{{cite book}}
: CS1 maint: location missing publisher (link)