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39-515: The Princeton Chert (Ashnola shale in older sources) and its fossils have been known since the 1950s, but have attracted increased attention in the late 1970 and on. This may be due to the rare type of silica permineralized fossil Lagerstätten found, which has preserved plants and animals in minute 3D detail, with exceptional internal cellular detail. This has meant anatomical descriptions and reconstruction of whole plants from isolated parts has been possible in many species. Few plant fossils elsewhere in

78-754: A Konservat-Lagerstätte may be based on a number of different factors which constitute "exceptional preservation". These may include the completeness of specimens, soft tissue preservation, fine-scale detail, taxonomic richness, distinctive taphonomic pathways (often multiple at the same site), the extent of the fossil layer in time and space, and particular sediment facies encouraging preservation. The world's major Lagerstätten include: Lakhanda Lagerstätte 1030-1000 Ma Uchur-Maya Depression, Russia Bitter Springs 1000–850 Ma South Australia Preserved fossils include cyanobacteria microfossils . Chichkan Lagerstätte 775 Ma Kazakhstan Doushantuo Formation 600–555 Ma Guizhou Province, China Spans

117-558: A death mask . It is a part of the larger Buen Formation , and has a fauna similar to the Maotianshan shales. Sinsk Algal Lens 518 Ma Yakutia , Russia One of the oldest known Cambrian lagerstätten . The fauna of this site is unique, as it seems that they were adapted to living in dysaerobic conditions. Pinus See List of Pinus species for complete taxonomy to species level. See list of pines by region for list of species by geographic distribution . A pine

156-400: A mycoparasite , Cryptodidymosphaerites princetonensis . Symbiotic mycorrhizal relationships have also been preserved in roots of Pinus and Metasequoia milleri . In Metasequoia these associations have been compared to extant mycorrhizae , and found to be very similar. The mycorrhizal relationship with Pinus was the first documentation of ectomycorrhizae from the fossil record, with

195-423: A freshwater fish, Amia (bowfin), have been found in the shale overlying the plant deposits, along with remains of the freshwater fishes Amyzon and Libotonius , plus a soft-shelled turtle. Pathogenic fungi have been recorded on the leaves and other organs of some vascular plants. Fossil Uhlia palms have tar spot fungi on their leaves described as Paleoserenomyces , which is in turn parasitized by

234-530: A huge ash fall, though few angiosperms occur in these layers. A large number of angiosperms have been found along with several types of conifers , ferns, and several unidentified fossils from various families. The array of floral and faunal fossils found in the Princeton Chert has offered unequivocal evidence that it was a lacustrine or lake environment. The plant fossils found show many structural and anatomical adaptations to an aquatic environment, including

273-564: A means to evaluate soil fertility and vigour of the trees. The bark of most pines is thick and scaly, but some species have thin, flaky bark. The branches are produced in "pseudo whorls", actually a very tight spiral but appearing like a ring of branches arising from the same point. Many pines are uninodal, producing just one such whorl of branches each year, from buds at the tip of the year's new shoot , but others are multinodal, producing two or more whorls of branches per year. Pines have four types of leaf : Pines are monoecious , having

312-487: A moist warm temperate climate with mean annual temperature 13.1 ± 3.1 °C, with mild winters (cold month mean temperature 5.3 ± 2.8 °C), and mean annual precipitation 114 ± 42 cm per year. Several of the smaller chert layers are separated by volcanic ash layers, indicating nearby volcanic activity. It is thought that fossils were pervaded with silicic acid due to this volcanic activity. Subsequently, water charged with minerals flowed from springs or geysers into

351-565: A reduced vascular system, aerenchyma in tissues (air spaces to provide buoyancy), and protoxylem lacunae surrounded by a ring of cells with thickened inner walls. Further evidence is provided by the fossils’ clear affinities with modern aquatic angiosperms. Many extant plants show these adaptations and are similar to the organisms found in the chert. For example, water lilies ( Allenbya , Nymphaeaceae ), water plantains ( Alismataceae ), arums ( Keratosperma , Araceae ) and rushes and sedges ( Ethela , Juncaceae / Cyperaceae ) are just some of

390-478: Is an interbedded sequence consisting of coal , shale , volcanic ash , and chert in the Allenby Formation . 49 chert layers, ranging in thickness from 1 to 55 cm (0.39 to 21.65 in) have been recognized and described, though each is not unique in organisms preserved. Despite this, trends are evident throughout the outcrop , with certain taxa appearing and disappearing with time. The Princeton Chert

429-530: Is any conifer tree or shrub in the genus Pinus ( / ˈ p aɪ n ə s / ) of the family Pinaceae . Pinus is the sole genus in the subfamily Pinoideae . World Flora Online accepts 187 species names of pines as current, with additional synonyms, making it the largest family among the conifers. The American Conifer Society (ACS) and the Royal Horticultural Society accept 121 species. The highest species diversity of pines

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468-777: Is found in Mexico. Pines are widely distributed in the Northern Hemisphere ; they occupy large areas of boreal forest , but are found in many habitats, including the Mediterranean Basin . The timber from pine trees is called "pine"; it is one of the most extensively used types of timber. There are currently 818 named cultivars (or trinomials ) recognized by the ACS. It is a well-known type of Christmas tree . Pine trees are evergreen , coniferous resinous trees (or, rarely, shrubs ) growing 3–80 metres (10–260 feet) tall, with

507-449: Is now restricted to fir ( Abies ) and Douglas-fir ( Pseudotsuga ). Pines are gymnosperms . The genus is divided into two subgenera based on the number of fibrovascular bundles in the needle. The subgenera can be distinguished by cone, seed, and leaf characters: Phylogenetic evidence indicates that both subgenera have a very ancient divergence from one another. Each subgenus is further divided into sections and subsections. Many of

546-785: Is one of the world's oldest living organisms at around 4,800 years old. This tree can be found in the White Mountains of California. An older tree, now cut down, was dated at 4,900 years old. It was discovered in a grove beneath Wheeler Peak and it is now known as " Prometheus " after the Greek immortal . The spiral growth of branches, needles, and cones scales are arranged in Fibonacci number ratios. The new spring shoots are sometimes called "candles"; they are covered in brown or whitish bud scales and point upward at first, then later turn green and spread outward. These "candles" offer foresters

585-629: Is the largest genus of the Pinaceae , the pine family, which first appeared in the Jurassic period. Based on recent Transcriptome analysis, Pinus is most closely related to the genus Cathaya , which in turn is closely related to spruces . These genera, with firs and larches , form the pinoid clade of the Pinaceae. Pines first appeared during the Early Cretaceous, with the oldest verified fossil of

624-941: The Devonian Hunsrück Slates and Gogo Formation , the Carboniferous Mazon Creek , the Triassic Madygen Formation , the Jurassic Posidonia Shale and Solnhofen Limestone , the Cretaceous Yixian , Santana , & Agua Nueva formations and the Tanis Fossil Site , the Eocene Fur Formation , Green River Formation , Messel Formation & Monte Bolca , the Miocene Foulden Maar and Ashfall Fossil Beds ,

663-507: The Himalayas and the Tibetan plateau due to the collision of tectonic plates. Also at the time, Australia , which was joined to Antarctica , was beginning to move northwards. The Princeton Chert fossils indicate that the area was an aquatic ecosystem , growing in tropical to subtropical conditions. More recent analysis of the fossil flora, however reconstructs for the Princeton Chert flora

702-578: The Pliocene Gray Fossil Site , and the Pleistocene Naracoorte Caves & La Brea Tar Pits . Palaeontologists distinguish two kinds: Konservat-Lagerstätten preserve lightly sclerotized and soft-bodied organisms or traces of organisms that are not otherwise preserved in the usual shelly and bony fossil record; thus, they offer more complete records of ancient biodiversity and behavior and enable some reconstruction of

741-584: The palaeoecology of ancient aquatic communities. In 1986, Simon Conway Morris calculated only about 14% of genera in the Burgess Shale had possessed biomineralized tissues in life. The affinities of the shelly elements of conodonts were mysterious until the associated soft tissues were discovered near Edinburgh, Scotland, in the Granton Lower Oil Shale of the Carboniferous . Information from

780-644: The Indo-European base *pīt- ‘resin’ (source of English pituitary ). Before the 19th century, pines were often referred to as firs (from Old Norse fura , by way of Middle English firre ). In some European languages, Germanic cognates of the Old Norse name are still in use for pines — in Danish fyr , in Norwegian fura/fure/furu , Swedish fura/furu , Dutch vuren , and German Föhre — but in modern English, fir

819-465: The Maotianshan shale the world's most important for understanding the evolution of early multi-cellular life. Qingjiang biota 518 Ma Hubei , China This site is particularly notable due to both the large proportion of new taxa represented (approximately 53% of the specimens), and the notable volume of soft-body tissue preservation. Sirius Passet 523-518 Ma Greenland A site known for its fauna, and that they were most likely preserved by

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858-419: The angiosperms found both today and in the Princeton Chert. Seeds have also been found which share adaptations with living aquatics. On the other hand, terrestrial fossils have rarely been found. The few that are, are represented mainly by seeds, some of which may have been transported by birds. Additional support for the aquatic nature of the Princeton Chert deposits comes from animal fossils. Several fossils of

897-462: The basin allowed peat to re-accumulate each time, producing the multiple layers. Sampling into the Princeton Chert has been carried out, but presently the data has not been analysed in detail. Across the outcrop, trends in taxa can be seen; in the topmost layers fossil organs of Metasequoia milleri cease to be represented, yet Pinus (pine) and monocotyledons increase in number. There is a huge increase in ferns , such as Dennstaedtiopsis , after

936-461: The broader range of organisms found in Lagerstätten have contributed to recent phylogenetic reconstructions of some major metazoan groups. Lagerstätten seem to be temporally autocorrelated, perhaps because global environmental factors such as climate might affect their deposition. A number of taphonomic pathways may produce Konservat-Lagerstätten : The identification of a fossil site as

975-670: The decomposition of both gross and fine biological features until long after a durable impression was created in the surrounding matrix. Fossil-Lagerstätten span geological time from the Neoproterozoic era to the present . Worldwide, some of the best examples of near-perfect fossilization are the Cambrian Maotianshan shales and Burgess Shale , the Ordovician Soom Shale , the Silurian Waukesha Biota ,

1014-993: The fungi suggested as close to the modern pine symbiotics Rhizopogon and Suillus Undescribed Undescribed Undescribed A lepidopteran forewing Not described to family/genus/species Undescribed Undescribed undescribed A turtle , Bones preserved in the Princeton Chert † Dickwhitea † Dickwhitea allenbyensis An athyriaceous fern † Makotopteris † Makotopteris princetonensis An athyriaceous fern † Trawetsia † Trawetsia princetonensis A blechnacious fern † Dennstaedtiopsis † Dennstaedtiopsis aerenchymata A dennstaedtioid fern Osmundaceae Osmunda Undescribed An osmundaceous fern Not identified to species Cupressaceae Metasequoia † Metasequoia milleri A dawn redwood † Pinus allisonii A 2-needled Pine † Pinus andersonii A 3-needled Pine † Pinus arnoldii A basal Pine , First described for ovulate cones A whole plant reconstruction includes

1053-550: The genus being Pinus yorkshirensis from the Hauterivian - Barremian boundary (~130-125 million years ago) from the Speeton Clay , England. The evolutionary history of the genus Pinus has been complicated by hybridization . Pines are prone to inter-specific breeding. Wind pollination, long life spans, overlapping generations, large population size, and weak reproductive isolation make breeding across species more likely. As

1092-488: The low lying basin where the Princeton chert was located. Here, the water surrounded organisms as they grew, along with plant debris which had been accumulated. Many organisms were preserved in situ, in the lake or small pond environment in which they lived. The preservation must have been rapid, due to the minute cellular detail which has been conserved. This sequence of events is thought to have been replicated up to 50 times, as

1131-570: The majority of species reaching 15–45 m (50–150 ft) tall. The smallest are Siberian dwarf pine and Potosi pinyon , and the tallest is an 83.45 m (273.8 ft) tall sugar pine located in Yosemite National Park . Pines are long lived and typically reach ages of 100–1,000 years, some even more. The longest-lived is the Great Basin bristlecone pine ( P. longaeva ). One individual of this species, dubbed " Methuselah ",

1170-507: The male and female cones on the same tree. The male cones are small, typically 1–5 cm long, and only present for a short period (usually in spring, though autumn in a few pines), falling as soon as they have shed their pollen . The female cones take 1.5–3 years (depending on species) to mature after pollination , with actual fertilization delayed one year. At maturity the female cones are 3–60 cm long. Each cone has numerous spirally arranged scales, with two seeds on each fertile scale;

1209-786: The poorly understood interval between the end of the Cryogenian period and the late Ediacaran Avalon explosion . Mistaken Point 565 Ma Newfoundland, Canada This site contains one of the most diverse and well-preserved collections of Precambrian fossils . Ediacara Hills 555 Ma South Australia The type location the Ediacaran period, and has preserved a significant amount of fossils from that time. Khatyspyt Lagerstätte 544 Ma Yakutia , Russia A Late Ediacaran lagerstätte preserving an Avalon-type biota. Maotianshan Shales (Chengjiang) 518 Ma Yunnan , China The preservation of an extremely diverse faunal assemblage renders

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1248-453: The scales at the base and tip of the cone are small and sterile, without seeds. The seeds are mostly small and winged, and are anemophilous (wind-dispersed), but some are larger and have only a vestigial wing, and are bird -dispersed. Female cones are woody and sometimes armed to protect developing seeds from foragers. At maturity, the cones usually open to release the seeds. In some of the bird-dispersed species, for example whitebark pine ,

1287-469: The seeds are only released by the bird breaking the cones open. In others, the seeds are stored in closed cones for many years until an environmental cue triggers the cones to open, releasing the seeds. This is called serotiny . The most common form of serotiny is pyriscence, in which a resin binds the cones shut until melted by a forest fire, for example in P. rigida . The modern English name "pine" derives from Latin pinus , which some have traced to

1326-422: The smaller groups of Pinus are composed of closely related species with recent divergence and history of hybridization. This results in low morphological and genetic differences. This, coupled with low sampling and underdeveloped genetic techniques, has made taxonomy difficult to determine. Recent research using large genetic datasets has clarified these relationships into the groupings we recognize today. Pinus

1365-675: The synonymized P. similkameenensis . † Pinus princetonensis A pinaceous cone Alismataceae † Heleophyton † Heleophyton helobieoides An aquatic or emergent water-plantain Lagerst%C3%A4tten A Fossil-Lagerstätte ( German: [ˈlaːɡɐˌʃtɛtə] , from Lager 'storage, lair' Stätte 'place'; plural Lagerstätten ) is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These formations may have resulted from carcass burial in an anoxic environment with minimal bacteria, thus delaying

1404-791: The world exhibit such excellence in both preservation and diversity. Similar aged fossil beds in Eocene lake sediments are found elsewhere in British Columbia , including in Driftwood Canyon Provincial Park near Smithers in northern British Columbia, the McAbee Fossil Beds west of Kamloops , about 160 km (160,000 m) NNW of the Princeton Chert beds, and the Klondike Mountain Formation around Republic, Washington , south of Princeton. The Princeton Chert

1443-450: Was originally considered to be Middle Eocene based on data from mammals , freshwater fish, and potassium-argon dates. Recently, more accurate radiometric techniques provided a date of 48.7 mya, placing the Princeton Chert in the Ypresian stage (47.8–56.0 mya), consistent with the whole Allenby Formation being now dated radiometrically as being Early Eocene . The climate at this time

1482-468: Was small. This long term warmth is thought to be due to increased greenhouse gases, particularly CO 2 trapping more heat. The reason for this sudden increase in CO 2 is unknown, but it is hypothesised that it was due to an increase in ocean floor being recycled via volcanic arcs and metamorphic decarbonation reactions. This happened because the ocean between India and Asia was disappearing and being replaced by

1521-637: Was warm; it had reached a maximum during a series of warming events during the Early Eocene with the Princeton Chert likely deposited after the Eocene Thermal Maximum 2 and during the Early Eocene Climatic Optimum. During this time the sea warmed approximately 4 °C and terrestrial temperatures were several degrees warmer than today, meaning little or no ice was present at the poles. The temperature difference between poles and equator

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