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91-505: Conifers are a group of cone-bearing seed plants , a subset of gymnosperms . Scientifically, they make up 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

182-603: 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

273-694: 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

364-446: A closed, damp cone of suitable size is inserted into a narrow-mouthed bottle and allowed to open upon drying. Cone cows are traditional homemade toys , made by children using material found in nature. The most common design is a spruce or pine cone with sticks or matches for legs, which can easily be attached by forcing them between the cone scales. Playing with cone cows often includes building an animal enclosure from sticks. For

455-509: A famous debate , which is said to exemplify the two major deviations in biological thinking at the time – whether animal structure was due to function or evolution. Most taxa differ morphologically from other taxa. Typically, closely related taxa differ much less than more distantly related ones, but there are exceptions to this. Cryptic species are species which look very similar, or perhaps even outwardly identical, but are reproductively isolated. Conversely, sometimes unrelated taxa acquire

546-422: 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

637-399: 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

728-437: 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 the total number of species is relatively small, conifers are ecologically important. They are the dominant plants over large areas of land, most notably

819-665: 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

910-589: A hard coat evolved to resist digestion in the bird's stomach. Members of the cypress family ( cypresses , arborvitae , junipers , redwoods , etc.) differ in that the bract and seed scales are fully fused, with the bract visible as no more than a small lump or spine on the scale. The botanical term galbulus (plural galbuli; from the Latin for a cypress cone) is sometimes used instead of strobilus for members of this family. The female cones have one to 20 ovules on each scale. They often have peltate scales, as opposed to

1001-413: 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

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1092-880: 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),

1183-402: 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

1274-536: A separate family (Taxodiaceae) in the past. In most of the genera, the cones are woody and the seeds have two narrow wings (one along each side of the seed), but in three genera ( Platycladus , Microbiota and Juniperus ), the seeds are wingless, and in Juniperus , the cones are fleshy and berry -like (known as galbuli ). The cones and seeds of Sciadopitys (the only member of the family) are similar to those of some Cupressaceae, but larger, 6–11 cm long;

1365-484: 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

1456-529: A similar appearance as a result of convergent evolution or even mimicry . In addition, there can be morphological differences within a species, such as in Apoica flavissima where queens are significantly smaller than workers. A further problem with relying on morphological data is that what may appear morphologically to be two distinct species may in fact be shown by DNA analysis to be a single species. The significance of these differences can be examined through

1547-410: A traditional part of the arts and crafts of cultures where conifers are common. Examples of their use includes seasonal wreaths and decorations, fire starters, bird feeders, toys, etc. An intriguing derivation of the impossible bottle mechanical puzzle takes advantage of the fact that pine cones open and close based on their level of dryness. In constructing the pine cone in the bottle display,

1638-468: 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 ,

1729-590: 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

1820-443: 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

1911-426: Is a fairground with cone cow sculptures large enough for children to ride on. In Sweden, a video game was released in which the player may build virtual cone cows. Swedish artist Lasse Åberg has created artwork with cone cows, which has been included in an alphabet book and featured on a Swedish postage stamp among other classic toys. Cones are also used as decorative elements in architecture such as on top of

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2002-414: 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

2093-495: Is an important indication of wildfire risk. Closed cones indicate damp conditions while open cones indicate the forest floor is dry. As a result of this, pine cones have often been used by people in temperate climates to predict dry and wet weather, usually hanging a harvested pine cone from some string outside to measure the humidity of the air. Members of the Araucariaceae ( Araucaria , Agathis , Wollemia ) have

2184-478: Is named after the pine cone. Pine cones were also used as symbols of fertility in ancient Assyrian art. In Christian symbolism, they are closely related to the tree of life . Morphology (biology) Morphology in biology is the study of the form and structure of organisms and their specific structural features. This includes aspects of the outward appearance (shape, structure, color, pattern, size), i.e. external morphology (or eidonomy ), as well as

2275-494: Is poisonous. The whole 'berry' with the seed is eaten by birds, which digest the sugar-rich scale and pass the hard seed undamaged in their droppings, so dispersing the seed far from the parent plant. Welwitschia is unique cone-bearing plant is not considered a conifer, but belongs in the Order Welwitschiales . Welwitschia mirabilis is often called a living fossil and is the only species in its genus, which

2366-483: Is that the male cones are located at the base of the branch, while the female at the tip (of the same or a different branchlet). However, in larches and cedars, both types of cones are always at the tips of short shoots, while both sexes of fir cones are always from side buds, never terminal. There is also some diversity in bearing in Cupressaceae. Some, Cupressus for instance, have little or no differentiation in

2457-432: 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

2548-598: Is the only genus in its family, which is the only family in its Order. The male cones are on male plants, and female cones on female plants. After emergence of the two cotyledons , it sets only two more leaves. Those two leaves then continue to grow longer from their base, much like fingernails. This allows it great drought tolerance, which is likely why it has survived in the desert of Namibia , while all other representatives from its order are now extinct. For most species found in Australia, male and female cones occur on

2639-583: 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

2730-741: 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

2821-413: The bract scales, and the seed scales (or ovuliferous scales), one subtended by each bract scale, derived from a highly modified branchlet . On the upper-side base of each seed scale are two ovules that develop into seeds after fertilization by pollen grains. The bract scales develop first, and are conspicuous at the time of pollination; the seed scales develop later to enclose and protect the seeds, with

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2912-460: 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

3003-484: 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,

3094-402: 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,

3185-408: 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

3276-436: 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 the one 11 m tall white spruce, Fraser et al. (1964) speculated that if the photosynthate used in making apical growth in 1961

3367-743: 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

3458-464: 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

3549-506: 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

3640-486: The apex of the cone are fertile, each bearing a single wingless seed, but in Saxegothaea several scales may be fertile. The fleshy scale complex is 0.5–3 cm long, and the seeds 4–10 mm long. In some genera (e.g. Prumnopitys ), the scales are minute and not fleshy, but the seed coat develops a fleshy layer instead, the cone having the appearance of one to three small plums on a central stem. The seeds have

3731-557: 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

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3822-435: The bract and seed scales fully fused, and have only one ovule on each scale. The cones are spherical or nearly so, and large to very large, 5–30 cm diameter, and mature in 18 months. For most species they disintegrate at maturity, to release the seeds although in some such as Araucaria bidwillii , the cone weighing up to 10 kilograms (22 lb) is shed intact. In Agathis , the seeds are winged and separate readily from

3913-458: The bract scales often not growing further. The scales open temporarily to receive pollen, then close during fertilization and maturation, and then re-open again at maturity to allow the seed to escape. Maturation takes 6–8 months from pollination in most Pinaceae genera, but 12 months in cedars and 18–24 months (rarely more) in most pines. The cones open either by the seed scales flexing back when they dry out, or (in firs, cedars and golden larch ) by

4004-530: 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 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

4095-552: The concept of form in biology, opposed to function , dates back to Aristotle (see Aristotle's biology ), the field of morphology was developed by Johann Wolfgang von Goethe (1790) and independently by the German anatomist and physiologist Karl Friedrich Burdach (1800). Among other important theorists of morphology are Lorenz Oken , Georges Cuvier , Étienne Geoffroy Saint-Hilaire , Richard Owen , Carl Gegenbaur and Ernst Haeckel . In 1830, Cuvier and Saint-Hilaire engaged in

4186-587: 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

4277-443: The cones disintegrating with the seed scales falling off. The cones are conic, cylindrical or ovoid (egg-shaped), and small to very large, from 2–60 cm long and 1–20 cm broad. After ripening, the opening of non- serotinous pine cones is associated with their moisture content—cones are open when dry and closed when wet. This assures that the small, windborne seeds will be dispersed during relatively dry weather, and thus,

4368-601: The counter-intuitive technique of "topping" and collection of cones from the severed crown tops at one-third the cost of collection from untopped trees and without decreasing cone production (Slayton 1969, Nienstaedt 1981). Most species of spruce are prone to the formation of pineapple gall pseudocones caused by the woolly aphids in the genus Adelges . These are not cones, although they closely resemble them. Alder ( Alnus ) trees are not conifers, but their mature seed bearing catkins closely resemble cones. Because of their widespread occurrence, conifer cones have been

4459-420: The different conifer families, and is often crucial for the identification of many species of conifers. The members of the pine family ( pines , spruces , firs , cedars , larches , etc.) have cones that are imbricate (that is, with scales overlapping each other like fish scales). These cones, especially the woody female cones, are considered the "archetypal" tree cones.The female cone has two types of scale:

4550-422: The distance traveled from the parent tree will be enhanced. A pine cone will go through many cycles of opening and closing during its life span, even after seed dispersal is complete. This process occurs with older cones while attached to branches and even after the older cones have fallen to the forest floor . The condition of fallen pine cones is a crude indication of the forest floor's moisture content, which

4641-613: 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

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4732-565: 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

4823-449: 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,

4914-466: 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

5005-492: 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

5096-567: The form and structure of internal parts like bones and organs , i.e. internal morphology (or anatomy ). This is in contrast to physiology , which deals primarily with function. Morphology is a branch of life science dealing with the study of the gross structure of an organism or taxon and its component parts. The etymology of the word "morphology" is from the Ancient Greek μορφή ( morphḗ ), meaning "form", and λόγος ( lógos ), meaning "word, study, research". While

5187-460: 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 a single white spruce tree from 1926 to 1961. Apical growth of

5278-419: 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,

5369-491: The imbricate cones described above, though some have imbricate scales. The cones are usually small, 0.3–6 cm or 1 ⁄ 8 – 2 + 3 ⁄ 8 inches long, and often spherical or nearly so, like those of Nootka cypress , while others, such as western redcedar and California incense-cedar , are narrow. The scales are arranged either spirally, or in decussate whorls of two (opposite pairs) or three, rarely four. The genera with spiral scale arrangement were often treated in

5460-530: 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

5551-535: 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

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5642-580: The latter used to flavor gin . Conifer cone The male cone ( microstrobilus or pollen cone ) is structurally similar across all conifers, differing only in small ways (mostly in scale arrangement) from species to species. Extending out from a central axis are microsporophylls (modified leaves). Under each microsporophyll is one or several microsporangia ( pollen sacs). The female cone ( megastrobilus , seed cone , or ovulate cone ) contains ovules which, when fertilized by pollen, become seeds. The female cone structure varies more markedly between

5733-417: 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

5824-513: 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, the Cephalotaxaceae may be better included within

5915-403: 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

6006-581: The most part, cone cows have been displaced by manufactured toys, at least in affluent countries, but the creation of cone cows still enjoys some popularity as an outdoor activity for children. Cone cows are a part of children's culture in Finland where they are known as Käpylehmä (plural: Käpylehmät ) and Sweden where they are known as kottkor or kottdjur (cone animals). Schools and other institutions teach children how to make cone cows as part of outdoors education on nature and history. In Finland there

6097-466: 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

6188-423: 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

6279-424: 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

6370-567: 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

6461-499: 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

6552-416: 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

6643-457: The positions of male and female cones. Cone crop potential can be predicted in various ways. An early indication of a potential crop can be a period of abnormally hot, dry weather at the time of bud differentiation, particularly if the current and preceding cone crops have been poor (Nienstaedt and Zasada 1990). Estimates of cone crop potential can be made by counting female reproductive buds in fall or winter, and an experienced observer can detect

6734-567: The posts surrounding Koper 's Da Ponte Fountain , the central element of the Fontana della Pigna in Rome or a bronze cone in the narthex of the Aachen Cathedral . Cones are also occasionally used as a charge in heraldic coats of arms. In some parts of Russia and Georgia , immature pine cones are harvested in late spring and boiled to make sweet preserves . The pineal gland

6825-512: 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

6916-452: 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

7007-579: The same plant ( tree or shrub ), with female usually on the higher branches towards the top of the plant. This distribution is thought to improve chances of cross-fertilization , as pollen is unlikely to be blown vertically upward within the crown of one plant, but can drift slowly upward in the wind, blowing from low on one plant to higher on another plant. In some conifers, male cones additionally often grow clustered in large numbers together, while female cones are more often produced singly or in only small clusters. A further characteristic arrangement of pines

7098-411: The scales are imbricate and spirally arranged, and have 5-9 ovules on each scale. Members of the yew family and the closely related Cephalotaxaceae have the most highly modified cones of any conifer. There is only one scale in the female cone, with a single poisonous ovule. The scale develops into a soft, brightly coloured sweet, juicy, berry-like aril which partly encloses the deadly seed. The seed alone

7189-493: The seed scale, but in the other two genera, the seed is wingless and fused to the scale. The cones of the Podocarpaceae are similar in function, though not in development, to those of the Taxaceae (q.v. below), being berry-like with the scales highly modified, evolved to attract birds into dispersing the seeds. In most of the genera, two to ten or more scales are fused together into a usually swollen, brightly coloured, soft, edible fleshy aril . Usually, only one or two scales at

7280-489: 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),

7371-427: 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

7462-404: 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 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

7553-437: 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 the years 1955 through 1961, respectively. The total number of needles of all ages present on

7644-475: The subtle morphological differences and distinguish between reproductive buds and vegetative buds (Eis 1967b). White spruce seed collection is expensive, and collection from cone caches of red squirrels is probably the cheapest method. The viability of seed from cached cones does not vary during current caching, but viability drops essentially to zero after being in caches for 1 or 2 years (Wagg 1964). Collection of cones in seed orchards has been facilitated by

7735-517: 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

7826-420: 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 is probably: first to apical growth and new needle formation, then to buds for the next year's growth, with

7917-474: 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

8008-576: The use of allometric engineering in which one or both species are manipulated to phenocopy the other species. A step relevant to the evaluation of morphology between traits/features within species, includes an assessment of the terms: homology and homoplasy . Homology between features indicates that those features have been derived from a common ancestor. Alternatively, homoplasy between features describes those that can resemble each other, but derive independently via parallel or convergent evolution . The invention and development of microscopy enabled

8099-416: 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

8190-464: 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

8281-473: 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 the photosynthate to produce energy to sustain respiration over this period, an amount estimated to be about 10% of

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