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Streptophyta

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Streptophyta ( / s t r ɛ p ˈ t ɒ f ɪ t ə , ˈ s t r ɛ p t oʊ f aɪ t ə / ), informally the streptophytes ( / ˈ s t r ɛ p t ə f aɪ t s / , from the Greek strepto 'twisted', for the morphology of the sperm of some members), is a clade of plants . The composition of the clade varies considerably between authors, but the definition employed here includes land plants and all green algae except the Chlorophyta and the more basal Prasinodermophyta .

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77-552: The composition of Streptophyta and similar groups (Streptophytina, Charophyta ) varies in each classification. Some authors include only the Charales and Embryophyta (e.g., Streptophyta, Streptophytina Lewis & McCourt 2004 ), others include more groups (e.g., Charophyta Lewis & McCourt 2004; Karol et al. 2009; Adl et al. 2012 , Streptophyta; de Reviers 2002; Leliaert et al. 2012, Streptobionta Kenrick & Crane 1997 ; some authors use this broader definition, but exclude

154-636: A division , yet also as a superdivision or an unranked clade . The terrestrial plants, the Embryophyta emerged deep within Charophyta, possibly from terrestrial unicellular charophytes, with the class Zygnematophyceae as a sister group . With the Embryophyta now cladistically placed in the Charophyte, it is a synonym of Streptophyta. The sister group of the charophytes are

231-604: A fossil record extending back about 300 million years to the Paleozoic in the late Carboniferous period; even many of the modern genera are recognizable from fossils 60–120 million years old. Other classes and orders, now long extinct, also occur as fossils, particularly from the late Paleozoic and Mesozoic eras. Fossil conifers included many diverse forms, the most dramatically distinct from modern conifers being some herbaceous conifers with no woody stems. Major fossil orders of conifers or conifer-like plants include

308-406: A gametophyte . In embryophytes (land plants) the zygote will instead give rise to a multicellular sporophyte . Except from land plants, retention of the zygote is only known from some species in one group of green algae; the coleochaetes . In these species the zygote is corticated by a layer of sterile gametophytic cells. Another similarity is the presence of sporopollenin in the inner wall of

385-697: A sister group to Pinales (the 'gnepine' hypothesis) or as being more derived than Pinales but sister to the rest of the group. Most recent studies favor the 'gnepine' hypothesis. The earliest conifers appear in the fossil record during the Late Carboniferous ( Pennsylvanian ), over 300 million years ago. Conifers are thought to be most closely related to the Cordaitales , a group of extinct Carboniferous-Permian trees and clambering plants whose reproductive structures had some similarities to those of conifers. The most primitive conifers belong to

462-1099: A basal Streptophyte. These earlier classifications have not taken into account that the Coleochaetophyceae and the Zygnemophyceae appear to have emerged in the Charophyceae + Embryophyta clade, resulting in the synonymy of the Phragmoplastophyta and Streptophytina/Streptophyta sensu stricto (a.k.a. Adl 2012) nomenclature. Below is a reconstruction of Streptophyta relationships, based on genomic data. Mesostigma viride (Mesostigmatophyceae s.s.) Spirotaenia Chlorokybophyceae Klebsormidiophyceae Charophyceae (Stoneworts & musk grasses) Coleochaetophyceae Zygnematophyceae (desmids, water silk etc.) Anthocerotophyta (hornworts) Marchantiophyta (Liverworts) Bryophyta (Mosses) † Horneophytopsida Tracheophyta Streptofilum , described in 2018, appears to bring in

539-422: A female multicellular gametophyte. The female gametophytes grow to produce two or more archegonia , each of which contains an egg. Upon fertilization, the diploid egg will give rise to the embryo, and a seed is produced. The female cone then opens, releasing the seeds which grow to a young seedling . Conifer reproduction is synchronous with seasonal changes in temperate zones. Reproductive development slows to

616-399: A female cone and are drawn into a tiny opening on the ovule called the micropyle . It is within the ovule that pollen-germination occurs. From here, a pollen tube seeks out the female gametophyte, which contains archegonia each with an egg, and if successful, fertilization occurs. The resulting zygote develops into an embryo , which along with the female gametophyte (nutritional material for

693-665: A halt during each winter season and then resumes each spring. The male strobilus development is completed in a single year. Conifers are classified by three reproductive cycles that refer to the completion of female strobilus development from initiation to seed maturation. All three types of reproductive cycle have a long gap between pollination and fertilization . One year reproductive cycle : The genera include Abies , Picea , Cedrus , Pseudotsuga , Tsuga , Keteleeria ( Pinaceae ) and Cupressus , Thuja , Cryptomeria , Cunninghamia and Sequoia ( Cupressaceae ) . Female strobili are initiated in late summer or fall of

770-413: A maximum of energy from weak sunshine at high latitudes or under forest canopy shade. Conifers from hotter areas with high sunlight levels (e.g. Turkish pine Pinus brutia ) often have yellower-green leaves, while others (e.g. blue spruce , Picea pungens ) may develop blue or silvery leaves to reflect ultraviolet light. In the great majority of genera the leaves are evergreen , usually remaining on

847-880: A needle-like appearance, but others, including most of the Cupressaceae and some of the Podocarpaceae , have flat, triangular scale-like leaves. Some, notably Agathis in Araucariaceae and Nageia in Podocarpaceae, have broad, flat strap-shaped leaves. Others such as Araucaria columnaris have leaves that are awl-shaped. In the majority of conifers, the leaves are arranged spirally, the exceptions being most of Cupressaceae and one genus in Podocarpaceae, where they are arranged in decussate opposite pairs or whorls of 3 (−4). In many species with spirally arranged leaves, such as Abies grandis (pictured),

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924-499: A new branching. Mesostigmatophyceae Chlorokybophyceae Streptofilum Klebsormidiophyceae Charophyceae (Stoneworts & musk grasses) Zygnematophyceae (desmids, water silk etc.) Coleochaetophyceae Embryophytes Charophyta Embryophyta Charophyta ( UK : / k ə ˈ r ɒ f ɪ t ə , ˌ k ær ə ˈ f aɪ t ə / ) is a group of freshwater green algae , called charophytes ( / ˈ k ær ə ˌ f aɪ t s / ), sometimes treated as

1001-402: A protective cone called a strobilus . The cones take from four months to three years to reach maturity, and vary in size from 2 to 600 millimetres ( 1 ⁄ 8 to 23 + 5 ⁄ 8  in) long. In Pinaceae , Araucariaceae , Sciadopityaceae and most Cupressaceae , the cones are woody , and when mature the scales usually spread open allowing the seeds to fall out and be dispersed by

1078-487: A serious environmental issue causing problems for pastoral farming and for conservation . Radiata pine was introduced to Australia in the 1870s. It is "the dominant tree species in the Australian plantation estate" – so much so that many Australians are concerned by the resulting loss of native wildlife habitat. The species is widely regarded as an environmental weed across southeastern and southwestern Australia and

1155-406: A single white spruce tree from 1926 to 1961. Apical growth of the stem was slow from 1926 through 1936 when the tree was competing with herbs and shrubs and probably shaded by larger trees. Lateral branches began to show reduced growth and some were no longer in evidence on the 36-year-old tree. Apical growth totaling about 340 m, 370 m, 420 m, 450 m, 500 m, 600 m, and 600 m was made by the tree in

1232-469: A useful guide by which to assess the adequacy of particular nutrients, and the ratios among the major nutrients are helpful guides to nutritional imbalances. The softwood derived from conifers is of great economic value, providing about 45% of the world's annual lumber production. Other uses of the timber include the production of paper and plastic from chemically treated wood pulp. Some conifers also provide foods such as pine nuts and juniper berries ,

1309-591: A year, then they overwinter. Female strobili emerge followed by pollination in the following spring. Fertilization takes place in summer of the following year, only 3–4 months after pollination. Cones mature and seeds are then shed by the end of that same year. Pollination and fertilization occur in a single growing season. Two-year reproductive cycle : The genera includes Widdringtonia , Sequoiadendron ( Cupressaceae ) and most species of Pinus . Female strobilus initials are formed in late summer or fall then overwinter. Female strobili emerge and receive pollen in

1386-446: Is a coast redwood ( Sequoia sempervirens ), with a height of 115.55 metres (although one mountain ash, Eucalyptus regnans , allegedly grew to a height of 140 metres, the tallest living angiosperms are significantly smaller at around 100 metres. ) The thickest (that is, the tree with the greatest trunk diameter ) is a Montezuma cypress ( Taxodium mucronatum ), 11.42 metres in diameter. The largest tree by three-dimensional volume

1463-418: Is a giant sequoia ( Sequoiadendron giganteum ), with a volume 1486.9 cubic metres. The smallest is the pygmy pine ( Lepidothamnus laxifolius ) of New Zealand, which is seldom taller than 30 cm when mature. The oldest non-clonal living tree is a Great Basin bristlecone pine ( Pinus longaeva ), 4,700 years old. Since most conifers are evergreens, the leaves of many conifers are long, thin and have

1540-417: Is a short, filamentous and unbranched algae surrounded by a mucilaginous sheath, which often disintegrates to diads and unicells. The cells in Charophyta algae are all haploid , except during sexual reproduction, where a diploid unicellular zygote is produced. The zygote becomes four new haploid cells through meiosis, which will develop into new algae. In multicellular forms these haploid cells will grow into

1617-402: Is probably: first to apical growth and new needle formation, then to buds for the next year's growth, with the cambium in the older parts of the branches receiving sustenance last. In the white spruce studied by Fraser et al. (1964), the needles constituted 17.5% of the over-day weight. Undoubtedly, the proportions change with time. Wind and animal dispersals are two major mechanisms involved in

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1694-432: Is the whitespotted sawyer ( Monochamus scutellatus ). Adults are found in summer on newly fallen or recently felled trees chewing tiny slits in the bark in which they lay eggs. The eggs hatch in about two weeks and the tiny larvae tunnel to the wood and score its surface with their feeding channels. With the onset of cooler weather, they bore into the wood, making oval entrance holes and tunnelling deeply. Feeding continues

1771-623: The Cephalotaxaceae may be better included within the Taxaceae, and some authors additionally recognize Phyllocladaceae as distinct from Podocarpaceae (in which it is included here). The family Taxodiaceae is here included in the family Cupressaceae, but was widely recognized in the past and can still be found in many field guides. A new classification and linear sequence based on molecular data can be found in an article by Christenhusz et al. The conifers are an ancient group, with

1848-554: The Chlorophyta . In some charophyte groups, such as the Zygnematophyceae or conjugating green algae, flagella are absent and sexual reproduction does not involve free-swimming flagellate sperm. Flagellate sperm, however, are found in stoneworts ( Charales ) and Coleochaetales , orders of parenchymatous charophytes that are the closest relatives of the land plants, where flagellate sperm are also present in all except

1925-583: The Cordaitales , Vojnovskyales , Voltziales and perhaps also the Czekanowskiales (possibly more closely related to the Ginkgophyta ). Pinaceae Araucariaceae Podocarpaceae Sciadopityaceae Cupressaceae Cephalotaxaceae Taxaceae Multiple studies also indicate that the Gnetophyta belong within the conifers despite their distinct appearances, either placing them as

2002-743: The Mesozoic era. Modern groups of conifers emerged from the Voltziales during the Late Permian through Jurassic . Conifers underwent a major decline in the Late Cretaceous corresponding to the explosive adaptive radiation of flowering plants . All living conifers are woody plants, and most are trees, the majority having a monopodial growth form (a single, straight trunk with side branches) with strong apical dominance . Many conifers have distinctly scented resin , secreted to protect

2079-599: The conifers and flowering plants . Fossil stoneworts of early Devonian age that are similar to those of the present day have been described from the Rhynie chert of Scotland. Somewhat different charophytes have also been collected from the Late Devonian (Famennian) Waterloo Farm lagerstätte of South Africa. These include two species each of Octochara and Hexachara , which are the oldest fossils of Charophyte axes bearing in situ oogonia . The name comes from

2156-678: The division Pinophyta ( / p ɪ ˈ n ɒ f ɪ t ə , ˈ p aɪ n oʊ f aɪ t ə / ), also known as Coniferophyta ( / ˌ k ɒ n ɪ f ə ˈ r ɒ f ɪ t ə , - oʊ f aɪ t ə / ) or Coniferae . The division contains a single extant class , Pinopsida . All extant conifers are perennial woody plants with secondary growth . The great majority are trees , though a few are shrubs . Examples include cedars , Douglas-firs , cypresses , firs , junipers , kauri , larches , pines , hemlocks , redwoods , spruces , and yews . As of 2002, Pinophyta contained seven families, 60 to 65 genera, and more than 600 living species. Although

2233-518: The genus Chara , but the finding that the Embryophyta actually emerged in them has not resulted in a much more restricted meaning of the Charophyta, namely to a much smaller side branch. This more restricted group corresponds to the Charophyceae . The Zygnematophyceae , formerly known as the Conjugatophyceae, generally possess two fairly elaborate chloroplasts in each cell, rather than many discoid ones. They reproduce asexually by

2310-460: The influence of environmental conditions, their anatomical characteristics record growth rate changes produced by these changing conditions. The microscopic structure of conifer wood consists of two types of cells : parenchyma , which have an oval or polyhedral shape with approximately identical dimensions in three directions, and strongly elongated tracheids. Tracheids make up more than 90% of timber volume. The tracheids of earlywood formed at

2387-402: The wind . In some (e.g. firs and cedars ), the cones disintegrate to release the seeds, and in others (e.g. the pines that produce pine nuts ) the nut-like seeds are dispersed by birds (mainly nutcrackers , and jays ), which break up the specially adapted softer cones. Ripe cones may remain on the plant for a varied amount of time before falling to the ground; in some fire-adapted pines,

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2464-409: The 16 nutrient elements known to be essential to plants, 13 of which are obtained from the soil, including nitrogen , phosphorus , potassium , calcium , magnesium , and sulfur , all used in relatively large amounts. Nutrient concentrations in conifers also vary with season, age, and kind of tissue sampled, and analytical technique. The ranges of concentrations occurring in well-grown plants provide

2541-406: The Embryophyta, e.g., Charophyta Leliaert et al. 2012, Charophyceae Mattox & Stewart, 1984 , Streptophycophytes de Reviers, 2002 ). The clade Streptophyta includes both unicellular and multicellular organisms. Streptophyta contains the freshwater charophyte green algae and all land plants that reproduce sexually by conjugation. Mesostigma viride , a unicellular green flagellate alga may be

2618-743: The North American Forest Tree Nursery Soils Workshop at Syracuse in 1980 provided strong contrary evidence: Bob Eastman, President of the Western Maine Forest Nursery Co. stated that for 15 years he has been successful in avoiding winter “burn” to Norway spruce and white spruce in his nursery operation by fertilizing with 50–80 lb/ac (56–90 kg/ha) nitrogen in September, whereas previously winter burn had been experienced annually, often severely. Eastman also stated that

2695-464: The Pinales without Taxales as paraphyletic , and the latter order is no longer considered distinct. A more accurate subdivision would be to split the class into three orders, Pinales containing only Pinaceae, Araucariales containing Araucariaceae and Podocarpaceae, and Cupressales containing the remaining families (including Taxaceae), but there has not been any significant support for such a split, with

2772-755: The West of Scotland, Eire, parts of Wales and of the Lake District. Klebsormidium , the type of the Klebsormidiophyceae , is a simple filamentous form with circular, plate-like chloroplasts, reproducing by fragmentation, by dorsiventral, biciliate swarmers and, according to Wille, a twentieth-century algologist, by aplanospores . Sexual reproduction is simple and isogamous (the male and female gametes are outwardly indistinguishable). The Charales (Charophyceae), or stoneworts, are freshwater and brackish algae with slender green or grey stems;

2849-478: The absence of phycobilins , the presence of chlorophyll a and chlorophyll b , cellulose in the cell wall and the use of starch , stored in the plastids, as a storage polysaccharide. The charophytes and embryophytes share several traits that distinguish them from the chlorophytes, such as the presence of certain enzymes (class I aldolase , Cu/Zn superoxide dismutase , glycolate oxidase , flagellar peroxidase ), lateral flagella (when present), and, in many species,

2926-631: The base of charophytes (streptophytes). The cladograms below show consensus phylogenetic relationships based on plastid genomes and a new proposal for a third phylum of green plants based on analysis of nuclear genomes. Chlorophyta Mesostigmatophyceae s.l. Klebsormidiophyceae Charophyceae Coleochaetophyceae Zygnematophyceae Embryophytes (land plants) Palmophyllophyceae Prasinodermophyceae Chlorophyta Mesostigmatophyceae s.l. Klebsormidiophyceae Charophyceae Coleochaetophyceae Zygnematophyceae Embryophytes (land plants) Mesostigmatophyceae s.l. in

3003-558: The beginning of a growing season have large radial sizes and smaller, thinner cell walls . Then, the first tracheids of the transition zone are formed, where the radial size of cells and the thickness of their cell walls changes considerably. Finally, latewood tracheids are formed, with small radial sizes and greater cell wall thickness. This is the basic pattern of the internal cell structure of conifer tree rings. Most conifers are monoecious , but some are subdioecious or dioecious ; all are wind-pollinated . Conifer seeds develop inside

3080-399: The cladograms corresponds to a clade of a narrower circumscription, Mesostigmatophyceae s.s., and a separate class Chlorokybophyceae, as used by AlgaeBase . The Mesostigmatophyceae are not filamentous, but the other basal charophytes (streptophytes) are. Pinophyta Conifers are a group of cone-bearing seed plants , a subset of gymnosperms . Scientifically, they make up

3157-587: The cone consists of several fused scales, while in others (e.g. Taxaceae), the cone is reduced to just one seed scale or (e.g. Cephalotaxaceae) the several scales of a cone develop into individual arils, giving the appearance of a cluster of berries. The male cones have structures called microsporangia that produce yellowish pollen through meiosis. Pollen is released and carried by the wind to female cones. Pollen grains from living pinophyte species produce pollen tubes, much like those of angiosperms. The gymnosperm male gametophytes (pollen grains) are carried by wind to

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3234-463: The corresponding organs in other algae. As a result of fertilization, a protonema is formed, from which the sexually reproducing algae develops. A new terrestrial genus found in sandy soil in the Czech Republic , Streptofilum , may belong in its own class due its unique phylogenetic position. A cell wall is absent, instead the cell membrane consists of many layers of specific scales. It

3311-569: The development of a septum between the two cell-halves or semi-cells (in unicellular forms, each daughter-cell develops the other semi-cell afresh) and sexually by conjugation, or the fusion of the entire cell-contents of the two conjugating cells. The saccoderm desmids and the placoderm or true desmids, unicellular or filamentous members of the Zygnematophyceae, are dominant in non-calcareous, acid waters of oligotrophic or primitive lakes (e.g. Wastwater), or in lochans, tarns and bogs, as in

3388-402: The dispersal of conifer seeds. Wind-born seed dispersal involves two processes, namely; local neighborhood dispersal and long-distance dispersal. Long-distance dispersal distances range from 11.9–33.7 kilometres (7.4–20.9 mi) from the source. Birds of the crow family, Corvidae , are the primary distributor of the conifer seeds. These birds are known to cache 32,000 pine seeds and transport

3465-567: The female cone is termed fruit , which undergoes ripening (maturation). It was found recently that the pollen of conifers transfers the mitochondrial organelles to the embryo , a sort of meiotic drive that perhaps explains why Pinus and other conifers are so productive, and perhaps also has bearing on observed sex-ratio bias. Conifers are heterosporous , generating two different types of spores: male microspores and female megaspores . These spores develop on separate male and female sporophylls on separate male and female cones. In

3542-449: The first year spring and become conelets. The conelet goes through another winter rest and, in the spring of the second year archegonia form in the conelet. Fertilization of the archegonia occurs by early summer of the second year, so the pollination-fertilization interval exceeds a year. After fertilization, the conelet is considered an immature cone. Maturation occurs by autumn of the second year, at which time seeds are shed. In summary,

3619-466: The following spring. Female strobili emerge then pollination occurs in spring of the second year then the pollinated strobili become conelets in the same year (i.e. the second year). The female gametophytes in the conelet develop so slowly that the megaspore does not go through free-nuclear divisions until autumn of the third year. The conelet then overwinters again in the free-nuclear female gametophyte stage. Fertilization takes place by early summer of

3696-492: The following summer when larvae occasionally return to the surface of the wood and extend the feeding channels generally in a U-shaped configuration. During this time, small piles of frass extruded by the larvae accumulate under logs. Early in the spring of the second year following egg-laying, the larvae, about 30 mm long, pupate in the tunnel enlargement just below the wood surface. The resulting adults chew their way out in early summer, leaving round exit holes, so completing

3773-399: The fourth year and seeds mature in the cones by autumn of the fourth year. The growth and form of a forest tree are the result of activity in the primary and secondary meristems , influenced by the distribution of photosynthate from its needles and the hormonal gradients controlled by the apical meristems. External factors also influence growth and form. Fraser recorded the development of

3850-426: The grey colour of many species results from the deposition of lime on the walls, masking the green colour of the chlorophyll. The main stems are slender and branch occasionally. Lateral branchlets occur in whorls at regular intervals up the stem, they are attached by rhizoids to the substrate. The reproductive organs consist of antheridia and oogonia , though the structures of these organs differ considerably from

3927-419: The growing embryo) and its surrounding integument, becomes a seed . Eventually, the seed may fall to the ground and, if conditions permit, grow into a new plant. In forestry , the terminology of flowering plants has commonly though inaccurately been applied to cone-bearing trees as well. The male cone and unfertilized female cone are called male flower and female flower , respectively. After fertilization,

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4004-530: The immense conifer forests of the world represent the largest terrestrial carbon sink . Conifers are of great economic value for softwood lumber and paper production. Conifer is a Latin word, a compound of conus (cone) and ferre (to bear), meaning "the one that bears (a) cone(s)". The division name Pinophyta conforms to the rules of the International Code of Nomenclature for algae, fungi, and plants (ICN), which state (Article 16.1) that

4081-535: The largest and economically most important component group of gymnosperms, but nevertheless they comprise only one of the four groups. The division Pinophyta consists of just one class, Pinopsida, which includes both living and fossil taxa. Subdivision of the living conifers into two or more orders has been proposed from time to time. The most commonly seen in the past was a split into two orders, Taxales (Taxaceae only) and Pinales (the rest), but recent research into DNA sequences suggests that this interpretation leaves

4158-417: The leaf bases are twisted to present the leaves in a very flat plane for maximum light capture. Leaf size varies from 2 mm in many scale-leaved species, up to 400 mm long in the needles of some pines (e.g. Apache pine, Pinus engelmannii ). The stomata are in lines or patches on the leaves and can be closed when it is very dry or cold. The leaves are often dark green in colour, which may help absorb

4235-402: The majority of opinion preferring retention of all the families within a single order Pinales, despite their antiquity and diverse morphology . There were seven families of conifers c.  2011 , with 65–70 genera and over 600 living species ( c.  2002 ). The seven most distinct families are linked in the box above right and phylogenetic diagram left. In other interpretations,

4312-403: The male cones, microspores are produced from microsporocytes by meiosis . The microspores develop into pollen grains, which contain the male gametophytes. Large amounts of pollen are released and carried by the wind. Some pollen grains will land on a female cone for pollination. The generative cell in the pollen grain divides into two haploid sperm cells by mitosis leading to the development of

4389-466: The names of higher taxa in plants (above the rank of family) are either formed from the name of an included family (usually the most common and/or representative), in this case Pinaceae (the pine family), or are descriptive. A descriptive name in widespread use for the conifers (at whatever rank is chosen) is Coniferae (Art 16 Ex 2). According to the ICN, it is possible to use a name formed by replacing

4466-486: The one 11 m tall white spruce, Fraser et al. (1964) speculated that if the photosynthate used in making apical growth in 1961 was manufactured the previous year, then the 4 million needles that were produced up to 1960 manufactured food for about 600,000 mm of apical growth or 730 g dry weight, over 12 million mm of wood for the 1961 annual ring, plus 1 million new needles, in addition to new tissue in branches, bark, and roots in 1960. Added to this would be

4543-424: The one-year and the two-year cycles differ mainly in the duration of the pollination-fertilization interval. Three-year reproductive cycle : Three of the conifer species are pine species ( Pinus pinea , Pinus leiophylla , Pinus torreyana ) which have pollination and fertilization events separated by a two-year interval. Female strobili initiated during late summer or autumn of a year, then overwinter until

4620-425: The overwintering storage capacity of stock thus treated was much improved (Eastman 1980). The concentrations of nutrients in plant tissues depend on many factors, including growing conditions. Interpretation of concentrations determined by analysis is easy only when a nutrient occurs in excessively low or occasionally excessively high concentration. Values are influenced by environmental factors and interactions among

4697-569: The paraphyletic assemblage of " walchian conifers ", which were small trees, and probably originated in dry upland habitats. The range of conifers expanded during the Early Permian ( Cisuralian ) to lowlands due to increasing aridity. Walchian conifers were gradually replaced by more advanced voltzialean or "transition" conifers. Conifers were largely unaffected by the Permian–Triassic extinction event , and were dominant land plants of

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4774-429: The photosynthate to produce energy to sustain respiration over this period, an amount estimated to be about 10% of the total annual photosynthate production of a young healthy tree. On this basis, one needle produced food for about 0.19 mg dry weight of apical growth, 3 mm wood, one-quarter of a new needle, plus an unknown amount of branch wood, bark and roots. The order of priority of photosynthate distribution

4851-561: The plant for several (2–40) years before falling, but five genera ( Larix , Pseudolarix , Glyptostrobus , Metasequoia and Taxodium ) are deciduous , shedding their leaves in autumn. The seedlings of many conifers, including most of the Cupressaceae, and Pinus in Pinaceae, have a distinct juvenile foliage period where the leaves are different, often markedly so, from the typical adult leaves. Tree rings are records of

4928-416: The pollen tube. At fertilization, one of the sperm cells unites its haploid nucleus with the haploid nucleus of an egg cell. The female cone develops two ovules, each of which contains haploid megaspores. A megasporocyte is divided by meiosis in each ovule. Each winged pollen grain is a four celled male gametophyte . Three of the four cells break down leaving only a single surviving cell which will develop into

5005-514: The removal of individual plants beyond plantations is encouraged. At least 20 species of roundheaded borers of the family Cerambycidae feed on the wood of spruce , fir , and hemlock (Rose and Lindquist 1985). Borers rarely bore tunnels in living trees, although when populations are high, adult beetles feed on tender twig bark, and may damage young living trees. One of the most common and widely distributed borer species in North America

5082-453: The same effect in 105-day-old white spruce. The general short-term effect of nitrogen fertilization on coniferous seedlings is to stimulate shoot growth more so than root growth (Armson and Carman 1961). Over a longer period, root growth is also stimulated. Many nursery managers were long reluctant to apply nitrogenous fertilizers late in the growing season, for fear of increased danger of frost damage to succulent tissues. A presentation at

5159-927: The seeds as far as 12–22 km (7.5–13.7 mi) from the source. The birds store the seeds in the soil at depths of 2–3 cm ( 3 ⁄ 4 – 1 + 1 ⁄ 4  in) under conditions which favor germination . Conifers are the dominant plants over large areas of land, most notably the taiga of the Northern Hemisphere , but also in similar cool climates in mountains further south. A number of conifers originally introduced for forestry have become invasive species in parts of New Zealand , including radiata pine ( Pinus radiata ), lodgepole pine ( P. contorta ), Douglas fir ( Pseudotsuga mensiezii ) and European larch ( Larix decidua ). In parts of South Africa , maritime pine ( Pinus pinaster ), patula pine ( P. patula ) and radiata pine have been declared invasive species. These wilding conifers are

5236-545: The seeds may be stored in closed cones for up to 60–80 years, being released only when a fire kills the parent tree. In the families Podocarpaceae , Cephalotaxaceae , Taxaceae , and one Cupressaceae genus ( Juniperus ), the scales are soft, fleshy, sweet, and brightly colored, and are eaten by fruit-eating birds, which then pass the seeds in their droppings. These fleshy scales are (except in Juniperus ) known as arils . In some of these conifers (e.g. most Podocarpaceae),

5313-429: The sole source of nitrogen guanidine compounds were less prominent. Durzan and Steward noted that their results, drawn from determinations made in late summer, did not rule out the occurrence of different interim responses at other times of the year. Ammonium nitrogen produced significantly heavier (dry weight) seedlings with a higher nitrogen content after 5 weeks than did the same amount of nitrate nitrogen. Swan found

5390-517: The termination -aceae in the name of an included family, in this case preferably Pinaceae , by the appropriate termination, in the case of this division -ophyta . Alternatively, " descriptive botanical names " may also be used at any rank above family. Both are allowed. This means that if conifers are considered a division, they may be called Pinophyta or Coniferae. As a class, they may be called Pinopsida or Coniferae. As an order they may be called Pinales or Coniferae or Coniferales . Conifers are

5467-570: The total number of species is relatively small, conifers are ecologically important. They are the dominant plants over large areas of land, most notably the taiga of the Northern Hemisphere , but also in similar cool climates in mountains further south. Boreal conifers have many wintertime adaptations. The narrow conical shape of northern conifers, and their downward-drooping limbs, help them shed snow. Many of them seasonally alter their biochemistry to make them more resistant to freezing. While tropical rainforests have more biodiversity and turnover,

5544-475: The tree against insect infestation and fungal infection of wounds. Fossilized resin hardens into amber , which has been commercially exploited historically (for example, in New Zealand's 19th-century kauri gum industry). The size of mature conifers varies from less than one metre to over 100 metres in height. The world's tallest, thickest, largest, and oldest living trees are all conifers. The tallest

5621-661: The use of phragmoplasts in mitosis . Thus Charophyta and Embryophyta together form the clade Streptophyta , excluding the Chlorophyta. Charophytes such as Palaeonitella cranii and possibly the yet unassigned Parka decipiens are present in the fossil record of the Devonian . Palaeonitella differed little from some present-day stoneworts. There is an emerging consensus on green algal relationships, mainly based on molecular data. The Mesostigmatophyceae (including Spirotaenia , and Chlorokybophyceae) are at

5698-416: The usual 2-year life cycle. Conifers – notably Abies (fir), Cedrus , Chamaecyparis lawsoniana (Lawson's cypress), Cupressus (cypress), juniper , Picea (spruce), Pinus (pine), Taxus (yew), Thuja (cedar) – have been the subject of selection for ornamental purposes. Plants with unusual growth habits, sizes, and colours are propagated and planted in parks and gardens throughout

5775-468: The world. Conifers can absorb nitrogen in either the ammonium (NH 4 ) or nitrate (NO 3 ) form, but the forms are not physiologically equivalent. Form of nitrogen affected both the total amount and relative composition of the soluble nitrogen in white spruce tissues (Durzan and Steward). Ammonium nitrogen was shown to foster arginine and amides and lead to a large increase of free guanidine compounds, whereas in leaves nourished by nitrate as

5852-403: The years 1955 through 1961, respectively. The total number of needles of all ages present on the 36-year-old tree in 1961 was 5.25 million weighing 14.25 kg. In 1961, needles as old as 13 years remained on the tree. The ash weight of needles increased progressively with age from about 4% in first-year needles in 1961 to about 8% in needles 10 years old. In discussing the data obtained from

5929-418: The zygote. In at least one species, it receives nourishment from the gametophyte through placental transfer cells. Charophyta are complex green algae that form a sister group to the Chlorophyta and within which the Embryophyta emerged. The chlorophyte and charophyte green algae and the embryophytes or land plants form a clade called the green plants or Viridiplantae , that is united among other things by

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