43-510: 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 (Kasimovian-Gzhelian; 299.0–306.5 Ma) vegetation of clastic soils and some peat soils. The best known callistophyte
86-734: A class Equisetopsida ( Embryophyta ) encompassing all land plants. This is referred to as Equisetopsida sensu lato to distinguish it from the narrower use to refer to horsetails alone, Equisetopsida sensu stricto . They placed the lycopods into subclass Lycopodiidae and the ferns, keeping the term monilophytes, into five subclasses, Equisetidae, Ophioglossidae, Psilotidae, Marattiidae and Polypodiidae, by dividing Smith's Psilotopsida into its two orders and elevating them to subclass (Ophioglossidae and Psilotidae). Christenhusz et al. (2011) followed this use of subclasses but recombined Smith's Psilotopsida as Ophioglossidae, giving four subclasses of ferns again. Christenhusz and Chase (2014) developed
129-634: A few species (e.g., Cyathea brownii on Norfolk Island and Cyathea medullaris in New Zealand ). Roots are underground non-photosynthetic structures that take up water and nutrients from soil . They are always fibrous and are structurally very similar to the roots of seed plants. As in all vascular plants , the sporophyte is the dominant phase or generation in the life cycle . The gametophytes of ferns, however, are very different from those of seed plants. They are free-living and resemble liverworts , whereas those of seed plants develop within
172-405: A group of vascular plants (plants with xylem and phloem ) that reproduce via spores and have neither seeds nor flowers . They differ from mosses by being vascular, i.e., having specialized tissues that conduct water and nutrients, and in having life cycles in which the branched sporophyte is the dominant phase. Ferns have complex leaves called megaphylls that are more complex than
215-487: A new classification of ferns and lycopods. They used the term Polypodiophyta for the ferns, subdivided like Smith et al. into four groups (shown with equivalents in the Smith system), with 21 families, approximately 212 genera and 10,535 species; This was a considerable reduction in the number of families from the 37 in the system of Smith et al., since the approach was more that of lumping rather than splitting. For instance
258-1415: A number of families were reduced to subfamilies. Subsequently, a consensus group was formed, the Pteridophyte Phylogeny Group (PPG), analogous to the Angiosperm Phylogeny Group , publishing their first complete classification in November 2016. They recognise ferns as a class, the Polypodiopsida, with four subclasses as described by Christenhusz and Chase, and which are phylogenetically related as in this cladogram: Equisetales Ophioglossales Psilotales Marattiales Osmundales Hymenophyllales Gleicheniales Schizaeales Vascular plant Vascular plants (from Latin vasculum 'duct'), also called tracheophytes ( UK : / ˈ t r æ k iː ə ˌ f aɪ t s / , US : / ˈ t r eɪ k iː ə ˌ f aɪ t s / ) or collectively tracheophyta ( / ˌ t r eɪ k iː ˈ ɒ f ɪ t ə / ; from Ancient Greek τραχεῖα ἀρτηρία ( trakheîa artēría ) 'windpipe' and φυτά ( phutá ) 'plants'), are plants that have lignified tissues (the xylem ) for conducting water and minerals throughout
301-462: A protective coating called an indusium . The arrangement of the sporangia is important in classification. In monomorphic ferns, the fertile and sterile leaves look morphologically the same, and both are able to photosynthesize. In hemidimorphic ferns, just a portion of the fertile leaf is different from the sterile leaves. In dimorphic (holomorphic) ferns, the two types of leaves are morphologically distinct . The fertile leaves are much narrower than
344-430: A short-lived structure anchored to the ground by rhizoids called gametophyte which produce gametes. When a mature fertile frond bears sori, and spores are released, the spores will settle on the soil and send out rhizoids , while it develops into a prothallus . The prothallus bears spherical antheridia ( s.g. antheridium ) which produce antherozoids (male gametophytes) and archegonia ( s.g. archegonium ) which release
387-568: A single oosphere . The antherozoid swims up the archegonium and fertilize the oosphere, resulting in a zygote, which will grow into a separate sporophyte, while the gametophyte shortly persists as a free-living plant. Carl Linnaeus (1753) originally recognized 15 genera of ferns and fern allies, classifying them in class Cryptogamia in two groups, Filices (e.g. Polypodium ) and Musci (mosses). By 1806 this had increased to 38 genera, and has progressively increased since ( see Schuettpelz et al (2018) ). Ferns were traditionally classified in
430-500: Is polyphyletic , the term fern allies should be abandoned, except in a historical context. More recent genetic studies demonstrated that the Lycopodiophyta are more distantly related to other vascular plants , having radiated evolutionarily at the base of the vascular plant clade , while both the whisk ferns and horsetails are as closely related to leptosporangiate ferns as the ophioglossoid ferns and Marattiaceae . In fact,
473-566: Is an antiquated remnant of the obsolete scala naturae , and the term is generally considered to be unscientific. Botanists define vascular plants by three primary characteristics: Cavalier-Smith (1998) treated the Tracheophyta as a phylum or botanical division encompassing two of these characteristics defined by the Latin phrase "facies diploida xylem et phloem instructa" (diploid phase with xylem and phloem). One possible mechanism for
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#1732797727100516-578: Is intermediate between the eusporangiate ferns and the leptosporangiate ferns. Rai and Graham (2010) broadly supported the primary groups, but queried their relationships, concluding that "at present perhaps the best that can be said about all relationships among the major lineages of monilophytes in current studies is that we do not understand them very well". Grewe et al. (2013) confirmed the inclusion of horsetails within ferns sensu lato , but also suggested that uncertainties remained in their precise placement. Other classifications have raised Ophioglossales to
559-484: Is one of the indexes to the base of what is called the Dicksonites plueckenetii Subzone, which marks a significant phase in the continent-wide changes to the wetland ("coal swamp") vegetation that took place in late Moscovian times in response to a combination of tectonics-induced landscape changes and climate change. The increase in abundance of the callistophytes coincided with a decline in abundance and diversity of
602-613: Is present, it is found in the stem. Their foliage may be deciduous or evergreen , and some are semi-evergreen depending on the climate. Like the sporophytes of seed plants, those of ferns consist of stems, leaves and roots. Ferns differ from spermatophytes in that they reproduce by spores rather than having flowers and producing seeds. However, they also differ from spore-producing bryophytes in that, like seed plants, they are polysporangiophytes , their sporophytes branching and producing many sporangia. Also unlike bryophytes, fern sporophytes are free-living and only briefly dependent on
645-479: The class Filices, and later in a Division of the Plant Kingdom named Pteridophyta or Filicophyta. Pteridophyta is no longer recognised as a valid taxon because it is paraphyletic . The ferns are also referred to as Polypodiophyta or, when treated as a subdivision of Tracheophyta (vascular plants), Polypodiopsida, although this name sometimes only refers to leptosporangiate ferns. Traditionally, all of
688-566: The microphylls of clubmosses . Most ferns are leptosporangiate ferns . They produce coiled fiddleheads that uncoil and expand into fronds . The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the Polypodiopsida , comprising both the leptosporangiate ( Polypodiidae ) and eusporangiate ferns , the latter group including horsetails , whisk ferns , marattioid ferns , and ophioglossoid ferns . The fern crown group , consisting of
731-422: The nucellus has a lagenostome -like projection, which breaks down to form the pollen chamber. The full ontogeny of these ovules has been worked out in some detail and seems to be essentially similar to that seen in modern-day gymnosperms , including the use of a pollen drop to help capture and draw the pollen into the pollen chamber and a pollen tube to deliver the generative nucleus. The ovules were borne on
774-508: The "true" tracheophytes, the eutracheophytes. † Aglaophyton † Horneophytopsida † Rhyniophyta Lycopodiophyta † Zosterophyllophyta † Cladoxylopsida Equisetopsida (horsetails) Marattiopsida Psilotopsida (whisk ferns and adders'-tongues) Pteridopsida (true ferns) † Progymnospermophyta Cycadophyta (cycads) Ginkgophyta (ginkgo) Gnetophyta Pinophyta (conifers) Magnoliophyta (flowering plants) † Pteridospermatophyta (seed ferns) This phylogeny
817-530: The Lyginopteridales, which occupied very similar ecological niches and were very similar in general habit. It seems possible, therefore, that the reproductively more sophisticated callistophytes were able to out-compete and replace the Lyginopteridales. The Callistophytaceae flourished in Euramerica through Late Pennsylvanian times, eventually becoming extinct as this part of Pangaea became arid at
860-399: The ability to grow independent roots, woody structure for support, and more branching. A proposed phylogeny of the vascular plants after Kenrick and Crane 1997 is as follows, with modification to the gymnosperms from Christenhusz et al. (2011a), Pteridophyta from Smith et al. and lycophytes and ferns by Christenhusz et al. (2011b) The cladogram distinguishes the rhyniophytes from
903-469: The characteristic secretory structures have an integument that was only fused to the nucellus in the basal part of the ovules and so superficially resemble medullosalean ovules . Unlike the Medullosales , the ovules appear to be bilaterally symmetrical, although details of the vasculature suggest they were in fact evolved from plants with radially symmetrical ovules . The apical part of
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#1732797727100946-926: The dichotomy. Such fronds, when found as adpressions, are known as Dicksonites . The pinnules tend to be tongue-shaped or somewhat lobed, and can closely resemble pinnules of Lyginopteridales fronds such as Mariopteris . Distinguishing the fronds of these two orders of pteridosperms can in fact be very difficult unless the pinnules are fertile, although generally the pinnule lamina of Dicksonites fronds tend to be somewhat vaulted, whereas Mariopteris pinnules are usually flatter. Occasional Vesicaspora pollen grains and fragments of Dicksonites -like fronds occur in early Moscovian (middle Westphalian) strata, especially in intra-montane basins. However, callistophyte foliage and pollen suddenly become relatively widespread and abundant in Euramerican floras in late Moscovian (late Asturian) floras. This increase in callistophyte abundance
989-431: The fronds are branched more than once, it can also be a combination of the pinnatifid are pinnate shapes. If the leaf blades are divided twice, the plant has bipinnate fronds, and tripinnate fronds if they branch three times, and all the way to tetra- and pentapinnate fronds. In tree ferns, the main stalk that connects the leaf to the stem (known as the stipe), often has multiple leaflets. The leafy structures that grow from
1032-494: The inclusion of Equisetaceae in the ferns, notably relating to the construction of their sperm and peculiarities of their roots. The leptosporangiate ferns are sometimes called "true ferns". This group includes most plants familiarly known as ferns. Modern research supports older ideas based on morphology that the Osmundaceae diverged early in the evolutionary history of the leptosporangiate ferns; in certain ways this family
1075-474: The leptosporangiate ferns. The Marattiaceae are a primitive group of tropical ferns with large, fleshy rhizomes and are now thought to be a sibling taxon to the leptosporangiate ferns. Several other groups of species were considered fern allies: the clubmosses , spikemosses , and quillworts in Lycopodiophyta ; the whisk ferns of Psilotaceae ; and the horsetails of Equisetaceae . Since this grouping
1118-585: The leptosporangiates and eusporangiates, is estimated to have originated in the late Silurian period 423.2 million years ago, but Polypodiales , the group that makes up 80% of living fern diversity, did not appear and diversify until the Cretaceous , contemporaneous with the rise of flowering plants that came to dominate the world's flora. Ferns are not of major economic importance, but some are used for food, medicine, as biofertilizer , as ornamental plants, and for remediating contaminated soil. They have been
1161-555: The maternal gametophyte . The green , photosynthetic part of the plant is technically a megaphyll and in ferns, it is often called a frond . New leaves typically expand by the unrolling of a tight spiral called a crozier or fiddlehead into fronds . This uncurling of the leaf is termed circinate vernation . Leaves are divided into two types: sporophylls and tropophylls. Sporophylls produce spores; tropophylls do not. Fern spores are borne in sporangia which are usually clustered to form sori . The sporangia may be covered with
1204-473: The plant. They also have a specialized non-lignified tissue (the phloem ) to conduct products of photosynthesis . The group includes most land plants ( c. 300,000 accepted known species) other than mosses . Vascular plants include the clubmosses , horsetails , ferns , gymnosperms (including conifers ), and angiosperms ( flowering plants ). They are contrasted with nonvascular plants such as mosses and green algae . Scientific names for
1247-421: The presumed evolution from emphasis on haploid generation to emphasis on diploid generation is the greater efficiency in spore dispersal with more complex diploid structures. Elaboration of the spore stalk enabled the production of more spores and the development of the ability to release them higher and to broadcast them further. Such developments may include more photosynthetic area for the spore-bearing structure,
1290-492: The purely vegetative fronds, and so can give a superficially similarity to fertile fern fronds. Unlike ferns, however, these pollen-organs produced monolete, bisaccate pollen (fossil genus Vesicaspora ) bearing some similarity to the pollen of many conifers. The foliage, which is the part of these plants most widely-found as macrofossils , consists of fronds with a basal dichotomy of the main rachis, each branch producing pinnately divided segments, but with no pinnae attached below
1333-430: The rank of a fifth class, separating the whisk ferns and ophioglossoid ferns. The ferns are related to other groups as shown in the following cladogram: Lycophytes [REDACTED] Ferns [REDACTED] Gymnosperms [REDACTED] Angiosperms [REDACTED] The classification of Smith et al. in 2006 treated ferns as four classes: In addition they defined 11 orders and 37 families. That system
Callistophytaceae - Misplaced Pages Continue
1376-476: The spore producing vascular plants were informally denominated the pteridophytes , rendering the term synonymous with ferns and fern allies . This can be confusing because members of the division Pteridophyta were also denominated pteridophytes ( sensu stricto ). Traditionally, three discrete groups have been denominated ferns: two groups of eusporangiate ferns, the families Ophioglossaceae ( adder's tongues , moonworts , and grape ferns) and Marattiaceae ; and
1419-417: The spore wall and are dependent on the parent sporophyte for their nutrition. A fern gametophyte typically consists of: The lifecycle of a fern involves two stages, as in club mosses and horsetails . In stage one, the spores are produced by sporophytes in sporangia , which are clustered together in sori ( s.g. sorus ), developing on the underside of fertile fronds. In stage two, the spores germinate into
1462-558: The start of Permian times. They extend through the Permian of China and include anatomically preserved ovules of Callospermarion , vegetative foliage of Emplecopteris triagularis , male organs of Norinotheca and ovulate fronds of Norinosperma, and stems of Calistophyton. Callistophytales appear to be victims of the Permo-Triassic extinction event in 'Cathaysia'. Fern The ferns ( Polypodiopsida or Polypodiophyta ) are
1505-470: The stems is the presence in the cortex of spherical secretory structures. Similar structures have also been found in associated ovules, pollen-organs and foliage, and were one of the main lines of evidence on which the reconstruction of the plant was based (compare with similar evidence used to reconstruct the Lyginopteris -bearing fossil plant). The small ovules (fossil genus Callospermarion ) with
1548-496: The sterile leaves, and may have no green tissue at all, as in the Blechnaceae and Lomariopsidaceae . The anatomy of fern leaves can be anywhere from simple to highly divided, or even indeterminate (e.g. Gleicheniaceae , Lygodiaceae ). The divided forms are pinnate , where the leaf segments are completely separated from one other, or pinnatifid (partially pinnate), where the leaf segments are still partially connected. When
1591-463: The stipe are known as pinnae and are often again divided into smaller pinnules. Fern stems are often loosely called rhizomes , even though they grow underground only in some of the species. Epiphytic species and many of the terrestrial ones have above-ground creeping stolons (e.g., Polypodiaceae ), and many groups have above-ground erect semi-woody trunks (e.g., Cyatheaceae , the scaly tree ferns). These can reach up to 20 meters (66 ft) tall in
1634-498: The subject of research for their ability to remove some chemical pollutants from the atmosphere. Some fern species, such as bracken ( Pteridium aquilinum ) and water fern ( Azolla filiculoides ), are significant weeds worldwide. Some fern genera, such as Azolla , can fix nitrogen and make a significant input to the nitrogen nutrition of rice paddies . They also play certain roles in folklore. Extant ferns are herbaceous perennials and most lack woody growth. When woody growth
1677-415: The underside of pinnules that did not differ significantly in form from those of the purely vegetative fronds. The pollen-producing organs (fossil genus Idanothekion ) consisted of small, radially symmetrical synangia, with each pollen-sac having a longitudinal dehisence structure. Like the ovules, the synangia were attached to the underside of pinnules that did not differ significantly in form from those of
1720-463: The vascular plants group include Tracheophyta, Tracheobionta and Equisetopsida sensu lato . Some early land plants (the rhyniophytes ) had less developed vascular tissue; the term eutracheophyte has been used for all other vascular plants, including all living ones. Historically, vascular plants were known as " higher plants ", as it was believed that they were further evolved than other plants due to being more complex organisms. However, this
1763-477: The whisk ferns and ophioglossoid ferns are demonstrably a clade , and the horsetails and Marattiaceae are arguably another clade. Smith et al. (2006) carried out the first higher-level pteridophyte classification published in the molecular phylogenetic era, and considered the ferns as monilophytes, as follows: Molecular data, which remain poorly constrained for many parts of the plants' phylogeny, have been supplemented by morphological observations supporting
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1806-844: Was a consensus of a number of studies, and was further refined. The phylogenetic relationships are shown in the following cladogram (to the level of orders). This division into four major clades was then confirmed using morphology alone. Lycopodiophytes (club mosses, spike mosses, quillworts) Spermatophytes (seed plants) Equisetales (horsetails) [REDACTED] Ophioglossales (grapeferns etc.) Psilotales (whisk ferns) [REDACTED] Marattiales [REDACTED] Osmundales [REDACTED] Hymenophyllales (filmy ferns) [REDACTED] Gleicheniales [REDACTED] Schizaeales Salviniales (heterosporous) Cyatheales (tree ferns) [REDACTED] Polypodiales [REDACTED] Subsequently, Chase and Reveal considered both lycopods and ferns as subclasses of
1849-472: Was documented from Late Pennsylvanian coal ball petrifactions in North America. The relatively slender stems (fossil genus Callistophyton ) had a eustele with a well-developed zone of secondary wood, and unlike most (but not all) other Palaeozoic pteridosperms, showed axillary branching. These characters strongly point to its having been a scrambling or climbing plant. A characteristic feature of
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