Pinus sylvestris

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Pinus sylvestris
Skuleskogen pine.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Pinophyta
Class: Pinopsida
Order: Pinales
Family: Pinaceae
Genus: Pinus
Subgenus: P. subg. Pinus
Section: P. sect. Pinus
Subsection: P. subsect. Pinus
Species:
P. sylvestris
Binomial name
Pinus sylvestris
L.
Pinus sylvestris range-01.png
Distribution

Pinus sylvestris, the Scots pine (UK), Scotch pine (US), Baltic pine, [2] or European red pine [3] is a species of tree in the pine family Pinaceae that is native to Eurasia. It can readily be identified by its combination of fairly short, blue-green leaves and orange-red bark.

Contents

Description

Young female cone Pinus sylvestris young female cone - Keila.jpg
Young female cone
Pinus sylvestris forest in Sierra de Guadarrama, central Spain Pinar Sierra de Guadarrama 2005-09-13.JPG
Pinus sylvestris forest in Sierra de Guadarrama, central Spain

Pinus sylvestris is an evergreen coniferous tree growing up to 35 metres (115 feet) in height [4] and 1 m (3 ft 3 in) in trunk diameter when mature, [5] exceptionally over 45 m (148 ft) tall and 1.7 m (5+1⁄2 ft) in trunk diameter on very productive sites. The tallest on record is a tree over 210 years old growing in Estonia which stands at 46.6 m (153 ft). [6] The lifespan is normally 150–300 years, with the oldest recorded specimens in Lapland, Northern Finland over 760 years. [7] [8] [9] [10]

The bark is thick, flaky and orange-red when young to scaly and gray-brown in maturity, sometimes retaining the former on the upper portion. [5] [9] [8] The habit of the mature tree is distinctive due to its long, bare and straight trunk topped by a rounded or flat-topped mass of foliage. [7] [8] [9] [10]

The shoots are light brown, with a spirally arranged scale-like pattern. On mature trees the leaves ('needles') are a glaucous blue-green, often darker green to dark yellow-green in winter, 2.5–5 centimetres (1–2 inches) long and 1–2 millimetres (1⁄32–3⁄32 in) broad, produced in fascicles of two with a persistent gray 5–10 mm (1⁄4–3⁄8 in) basal sheath. On vigorous young trees the leaves can be twice as long, and occasionally occur in fascicles of three or four on the tips of strong shoots. Leaf persistence varies from two to four years in warmer climates, and up to nine years in subarctic regions. Seedlings up to one year old bear juvenile leaves; these are single (not in pairs), 2–3 cm (3⁄4–1+1⁄4 in) long, flattened, with a serrated margin. [7] [9] [10]

Mature open cones and seeds Pinus sylvestris MHNT.BOT.2005.0.971.jpg
Mature open cones and seeds
Roots of an old pine in Ystad, Sweden Rotter - Ystad-2020.jpg
Roots of an old pine in Ystad, Sweden

The seed cones are red at pollination, then pale brown, globose and 4–8 mm (5⁄32–5⁄16 in) in diameter in their first year, expanding to full size in their second year, pointed ovoid-conic, green, then gray-green to yellow-brown at maturity, 3–7.5 cm (1+1⁄8–3 in) long. The cone scales have a flat to pyramidal apophysis (the external part of the cone scale), with a small prickle on the umbo (central boss or protuberance). The seeds are blackish, 3–5 mm (1⁄8–3⁄16 in) in length with a pale brown 12–20 mm (1⁄2–13⁄16 in) wing and are released when the cones open in spring 22–24 months after pollination. The pollen cones are yellow, occasionally pink, 8–12 mm (5⁄16–15⁄32 in) long; pollen release is in mid to late spring. [7] [9]

Varieties

Pinus sylvestris var. hamata, Crimea Pinus sylvestris hamata Babugan.jpg
Pinus sylvestris var. hamata, Crimea

Over 100 Pinus sylvestris varieties have been described in the botanical literature, but only three or four are now accepted. [11] They differ only minimally in morphology, but with more pronounced differences in genetic analysis and resin composition. Populations in westernmost Scotland are genetically distinct from those in the rest of Scotland and northern Europe, but not sufficiently to have been distinguished as separate botanical varieties. Trees in the far north of the range were formerly sometimes treated as var. lapponica, but the differences are clinal and it is not genetically distinct. [7] [8] [12] [13] [14] [15] [16] [17] [18] [19] [ excessive citations ]

ImageVarietiesDescriptionDistribution
Conifere Dordogne.jpg Pinus sylvestris var. sylvestris L., 1753Described above.The bulk of the range, from Scotland and Spain to central Siberia.
Pinus sylvestris hamata cones Bulgaria.jpg Pinus sylvestris var. hamataStevenFoliage more consistently glaucous all year, not becoming duller in winter; cones more frequently with a pyramidal apophysis.The Balkans, northern Turkey, Crimea, and the Caucasus.
Pinus sylvestris var mongolica kz01.jpg Pinus sylvestris var. mongolicaLitv.Foliage duller green, shoots gray-green; leaves occasionally up to 12 cm long.Mongolia and adjoining parts of southern Siberia and northwestern China.
Pinus-sylvestris-cone-2.jpg Pinus sylvestris var. nevadensisD.H.Christ.(not considered distinct from var. sylvestris by all authors) Kalenicz. Ex Kom. Cones often with thicker scales, but doubtfully distinguishable on morphology.The Sierra Nevada in southern Spain and possibly other Spanish populations
Pinus sylvestris 2020 G2.jpg Pinus sylvestris var. cretaceaKalenicz. ex Kom.From border regions between Russia and Ukraine. [20]

Names

Before the 18th century, the species was more often known as Scots fir or Scotch fir. Another, less common name is European redwood. [2]

The timber from it is also called red deal [21] or yellow deal; the name "deal" comes from an archaic unit of volume used to measure wood.

Distribution and habitat

Scots Pin Uto, January 2013.jpg
Scots pine in Stockholm archipelago, Sweden

Pinus sylvestris is the only pine native to northern Europe, [22] [23] ranging from Western Europe to Eastern Siberia, south to the Caucasus Mountains and Anatolia, and north to well inside the Arctic Circle in Fennoscandia. In the north of its range, it occurs from sea level to 1,000 m (3,300 ft), while in the south of its range it is a mountain tree, growing at 1,200–2,600 m (3,900–8,500 ft) altitude. [7] [8] [9] [12] Its distribution intersects with T. piniperda's habitat, making the beetle a primary pest of the tree.

The species is mainly found on poorer, sandy soils, rocky outcrops, peat bogs or close to the forest limit. On fertile sites, the pine is out-competed by other tree species, usually spruce or broad-leaved trees. [24]

Britain and Ireland

Scattered survivors (two recently dead) of extensive deforestation at Glen Quoich, Scotland Scots Pines Beinn a' Bhuird.jpg
Scattered survivors (two recently dead) of extensive deforestation at Glen Quoich, Scotland

The tree spread across Britain and Ireland after the Last Glacial Maximum. Pollen records show that pine was present locally in southern England by 9,000 years ago having entered from northeast France and that it had spread as far north as the Lake District and North Pennines 500 years later.

It was present in Ireland over 8,800 years ago but absent from Wales at that time which suggests that the pine in Ireland had a separate Iberian origin or contained surviving populations, although evidence towards its survival is lacking. [15] Pine expanded into Scotland between 8,000 and 8,500 years ago either from an independent refuge, from Scandinavia (via Doggerland) or from Ireland. As the climate warmed it became extinct from most of Britain and Ireland around 5,500 years ago except in Scotland, Kielder in England and The Burren in County Clare, Ireland.

The Irish and western Scottish populations went through a massive decline around 4,000 years ago which ultimately led to the practical extinction of the Irish population between 2,000 and 1,000 years ago. It was replaced by large areas of blanket bog in western Scotland and Ireland though the reasons for its decline and extinction in England are not clear, but it may have been influenced by human activities. [25]

In Britain it now occurs naturally only in Scotland. Historical and archaeological records indicate that it also occurred in Wales and England until about 300–400 years ago, becoming extinct there due to over-exploitation and grazing; it has been re-introduced in these countries. Similar historical extinction and re-introduction applies to Ireland, Denmark and the Netherlands. [9] [12] [10] [26] Whether it truly became extinct in England is unknown. It has been speculated that it may have survived wild long enough for trees used in cultivation in England to derive from native (rather than imported) sources. [27] Shakespeare (in Richard II ) was familiar with the species in the 1590s, as was Evelyn in the early 1660s ( Sylva ), both around the time when the pine was thought to become extinct in England, but when landowners were also beginning ornamental and forestry planting. [27]

The pine formed much of the Caledonian Forest, which once covered much of the Scottish Highlands. Overcutting for timber demand, fire, overgrazing by sheep and deer, and even deliberate clearance to deter wolves have all been factors in the decline of this once great pine and birch forest. Only comparatively small areas –17,000 hectares (42,000 acres), only just over 1% of the estimated original 1,500,000 ha (3,700,000 acres) [28]  –of this ancient forest remain, the main surviving remnants being at Abernethy Forest, Glen Affric, Rothiemurchus Forest, and the Black Wood of Rannoch. Plans are currently in progress to restore at least some areas and work has started at key sites. [9] [10]

Ecology

It forms either pure forests or mixes with Norway spruce, common juniper, silver birch, European rowan, Eurasian aspen and other hardwood species. In central and southern Europe, it occurs with numerous additional species, including European black pine, mountain pine, Macedonian pine, and Swiss pine. In the eastern part of its range, it occurs with Siberian pine, among others. [8] [9]

In 2020, black spot needle blight was found on hundreds of Pinus sylvestris var. mongolica trees in four forest farms in northeastern China. It first appeared on the upper part of the needles, and then the needles became withered and gradually showed light black spots, although they still remained green. As the fungal disease progressed, the needles eventually died and turned gray with many dark black spots. The fungus was identified as Heterotruncatella spartii (within the family Sporocadaceae) based on morphology and molecular methods. [29]

Uses

Botanical plate Pinus sylvestris Sturm01.jpg
Botanical plate
Scots pine forest in Estonia Meenikunno maastikukaiteala.jpg
Scots pine forest in Estonia

Pinus sylvestris is an important tree in forestry. The wood is used for pulp and sawn timber products. A seedling stand can be created by planting, sowing, or natural regeneration. Commercial plantation rotations vary between 50 and 120 years, with longer rotations in northeastern areas where growth is slower.

In Scandinavian countries, the pine was used for making tar in the preindustrial age. Some active tar producers still exist, but that industry has almost ceased. [10] [13] The pine has also been used as a source of rosin and turpentine.

The wood is pale brown to red-brown, and used for general construction work. It has a dry density around 470 kg/m3 (varying with growth conditions), an open porosity of 60%, a fibre saturation point of 0.25 kg/kg, and a saturation moisture content of 1.60 kg/kg. [13] The pine fibres are used to make the textile known as vegetable flannel, [30] which has a hemp-like appearance, but with a tighter, softer texture. [31]

The pine has also been widely planted in New Zealand and much of the colder regions of North America; it was one of the first trees introduced to North America, in about 1600. [32] It is listed as an invasive species in some areas there, including Ontario, [33] Michigan. [34] It has been widely used in the United States for the Christmas tree trade, and was one of the most popular Christmas trees from the 1950s through the 1980s. It remains popular for that usage, though it has been eclipsed in popularity, by such species as Fraser fir, Douglas-fir, and others. Despite its invasiveness in parts of eastern North America, the pine does not often grow well there, partly due to climate and soil differences between its native habitat and that of North America, and partly due to damage by pests and diseases; the tree often grows in a twisted, haphazard manner if not tended to (as they are in the Christmas tree trade). [8] [22] The pines may be killed by the pine wood nematode, which causes pine wilt disease. The nematode most often attacks trees that are at least ten years old and often kills trees it infects within a few weeks. [35]

Previously, the pine was grown in and used extensively by the coal mining regions of Flanders, Belgium. It was used to fortify tunnels, primarily because it would make a cracking sound when in need of replacement. Large patches of forest, mostly containing the species, are still scattered over the countryside.

Cultivars

Several cultivars are grown for ornamental purposes in parks and large gardens, of which 'Aurea', [36] 'Beuvronensis', [37] 'Frensham', [38] and 'Gold Coin' [39] have gained the Royal Horticultural Society's Award of Garden Merit. [40]

In culture

The Scots pine is the plant badge of Clan Gregor. It is the national tree of Scotland. [41]

Fossil record

One fossil seed cone of Pinus montana fossilis was sent by the Naturmuseum Senckenberg to the Swedish Museum of Natural History (Swedish: Naturhistoriska Riksmuseet), as a scientific gift specimen, the seed cone is of late Pliocene age (Reuverian). Pinus montana is a synonym of Pinus sylvestris. The cone fossil had been recovered during the years 1884 and 1885 in Niederrad which is a quarter of Frankfurt am Main, Germany. [42] [43]

Selection in haploid versus diploid tissue

Genes of Scots Pine that are expressed in the haploid stage of the life cycle appear to be subject to stronger purifying selection than genes expressed only in the diploid stage. [44] The concept that those genes of an organism that are expressed in the haploid stage are subject to more efficient natural selection than those genes expressed exclusively in the diploid stage is referred to as the “masking theory”. [44] This theory implies that purifying selection is more efficient in the haploid stage of the life cycle where fitness effects are more evidently expressed than in the diploid stage of the life cycle.

See also

Related Research Articles

<span class="mw-page-title-main">Pine</span> Genus of plants in the conifer family Pinaceae

A pine is any conifer tree or shrub in the genus Pinus of the family Pinaceae. Pinus is the sole genus in the subfamily Pinoideae.

<span class="mw-page-title-main">Douglas fir</span> Species of tree

The Douglas fir is an evergreen conifer species in the pine family, Pinaceae. It is native to western North America and is also known as Douglas-fir, Douglas spruce, Oregon pine, and Columbian pine. There are three varieties: coast Douglas-fir, Rocky Mountain Douglas-fir and Mexican Douglas-fir.

<span class="mw-page-title-main">Conifer</span> Group of cone-bearing seed plants

Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta, also known as Coniferophyta 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.

<span class="mw-page-title-main">Bristlecone pine</span> Three species of pine trees native to the Western United States

The term bristlecone pine covers three species of pine tree. All three species are long-lived and highly resilient to harsh weather and bad soils. One of the three species, Pinus longaeva, is among the longest-lived life forms on Earth. The oldest of this species is more than 4,800 years old, making it the oldest known individual of any species. Many scientists are curious as to why this tree is able to live so long. In one study, they discovered that Pinus longaeva has higher levels of telomerase activity, which further slows or prevents the attrition rate of telomeres. This potentially contributes to the extended life of the bristlecone pine.

<i>Pinus taeda</i> Species of conifer

Pinus taeda, commonly known as loblolly pine, is one of several pines native to the Southeastern United States, from East Texas to Florida, and north to southern New Jersey. The wood industry classifies the species as a southern yellow pine. U.S. Forest Service surveys found that loblolly pine is the second-most common species of tree in the United States, after red maple. For its timber, the pine species is regarded as the most commercially important tree in the Southeastern U.S. The common name loblolly is given because the pine species is found mostly in lowlands and swampy areas.

<i>Pinus mugo</i> Species of plant

Pinus mugo, known as dwarf mountain pine, mountain pine, scrub mountain pine, Swiss mountain pine, bog pine, creeping pine, or mugo pine, is a species of conifer, native to high elevation habitats from southwestern to Central Europe and Southeast Europe.

<i>Pinus nigra</i> Species of conifer

Pinus nigra, the Austrian pine or black pine, is a moderately variable species of pine, occurring across Southern Europe from the Iberian Peninsula to the eastern Mediterranean, on the Anatolian peninsula of Turkey, Corsica and Cyprus, as well as Crimea and in the high mountains of Northwest Africa. The world's oldest black pine, located in the Banaz district of Uşak, Türkiye, is estimated to be 1000 years old and is the oldest larch in the world. It has a length of 11 meters, a diameter of 3 meters and a circumference of 9.60 meters.

<i>Pinus jeffreyi</i> Pine tree found in North America

Pinus jeffreyi, also known as Jeffrey pine, Jeffrey's pine, yellow pine and black pine, is a North American pine tree. It is mainly found in California, but also in the westernmost part of Nevada, southwestern Oregon, and northern Baja California. It is named in honor of its botanist documenter John Jeffrey.

<i>Picea abies</i> Species of plant

Picea abies, the Norway spruce or European spruce, is a species of spruce native to Northern, Central and Eastern Europe.

<i>Pinus contorta</i> Species of plant

Pinus contorta, with the common names lodgepole pine and shore pine, and also known as twisted pine, and contorta pine, is a common tree in western North America. It is common near the ocean shore and in dry montane forests to the subalpine, but is rare in lowland rain forests. Like all pines, it is an evergreen conifer.

<i>Pinus resinosa</i> Species of plant (coniferous tree)

Pinus resinosa, known as red pine, is a pine native to Eastern North America.

<i>Pinus wallichiana</i> Species of conifer

Pinus wallichiana is a coniferous evergreen tree native to the Himalaya, Karakoram and Hindu Kush mountains, from eastern Afghanistan east across northern Pakistan and north west India to Yunnan in southwest China. It grows in mountain valleys at altitudes of 1800–4300 m, reaching 30–50 m (98–164 ft) in height. It favours a temperate climate with dry winters and wet summers. In Pashto, it is known as Nishtar.

<i>Pinus densiflora</i> Species of conifer

Pinus densiflora, also called the Japanese red pine, the Japanese pine, or Korean red pine, is a species of pine tree native to East Asia and Siberia. In China, the plant is known as 赤松.

<span class="mw-page-title-main">Coulter pine</span> Pine tree found in North America

Coulter pine, or big-cone pine, is a native of the coastal mountains of Southern California in the United States and northern Baja California in Mexico. Isolated groves are found as far north as Clearlake, California on the flanks of Mt. Konocti and Black Diamond Mines Regional Preserve. It is named after Thomas Coulter, an Irish botanist and physician. The Coulter pine produces some of the heaviest cones of any pine tree, up to 5 kg (11 lb) and among conifers is exceeded only by the cones of Araucaria bidwillii.

<span class="mw-page-title-main">Stone pine</span> Species of pine tree

The stone pine, botanical name Pinus pinea, also known as the Italian stone pine, Mediterranean stone pine, umbrella pine and parasol pine, is a tree from the pine family (Pinaceae). The tree is native to the Mediterranean region, occurring in Southern Europe and the Levant. The species was introduced into North Africa millennia ago, and is also naturalized in the Canary Islands, South Africa and New South Wales.

<i>Pinus cembra</i> Species of plant

Pinus cembra, also known as Swiss pine, Swiss stone pine, Arolla pine, Austrian stone pine, or just stone pine, is a species of pine tree in the subgenus Strobus.

<i>Pinus heldreichii</i> Species of conifer

Pinus heldreichii, the Bosnian pine or Heldreich's pine, is a species of pine native to mountainous areas of the Balkans and southern Italy.

<i>Pinus halepensis</i> Species of conifer

Pinus halepensis, commonly known as the Aleppo pine, also known as the Jerusalem pine, is a pine native to the Mediterranean region. It was officially named by the botanist Philip Miller in his 1768 book The Gardener's Dictionary; he probably never went to Aleppo but mentions seeing large specimens at Goodwood in the garden of the Duke of Richmond, which were transplanted in 1739.

<i>Pinus monophylla</i> Pine tree found in North America

Pinus monophylla, the single-leaf pinyon, is a pine in the pinyon pine group, native to North America. The range is in southernmost Idaho, western Utah, Arizona, southwest New Mexico, Nevada, eastern and southern California and northern Baja California.

<i>Pinus <span style="font-style:normal;">×</span> rhaetica</i> Nothospecies of plant

Pinus × rhaetica, commonly known as Rhætic pine, is a natural hybrid of mountain pine and Scots pine. It is a coniferous tree nothospecies located in the subsection Pinus. The species name refers to the Rhætica alps, located in eastern Switzerland and western Austria.

References

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  3. ↑ Wu, Fan; Sun, Xiaobo; Hu, Xingfeng; Zou, Bingzhang; Lin, Nengqing; Lin, Jingquan; Ji, Kongshu (2020). "Response of Nitrogen Metabolism in Masson Pine Needles to Elevated CO2". Forests. 11 (4): 390. doi: 10.3390/f11040390 .
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  6. ↑ Oidermaa, J.-J., ed. (9 February 2016). "Eesti kõrgeim mänd osutus hiiglaseks ka ülejäänud maailmas" [Estonia's tallest pine turned out to be a giant in the rest of the world as well]. ERR Novaator (in Estonian). Tallinn: Eesti Rahvusringhääling. Retrieved 9 February 2016.
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  21. ↑ "Red Deal/European Red Wood". Tralee: Adanack. Retrieved 24 October 2022.
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  24. ↑ Mátyás, C.; Ackzell, L.; Samuel, C. J. A. (2004). Scots pine – Pinus sylvestris (PDF). EUFORGEN Technical Guidelines for Genetic Conservation and Use. Rome: International Plant Genetic Resources Institute. p. 6. ISBN   978-92-9043-661-4. Archived from the original (PDF) on 19 January 2017. Retrieved 20 May 2023.
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  26. ↑ Carlisle, A.; Brown, A. H. F. (1968). "Biological Flora of the British Isles: Pinus sylvestris L.". Journal of Ecology. 56 (1): 269–307. doi:10.2307/2258078. JSTOR   2258078.
  27. 1 2 Edlin, H. L. (1970). Trees, Woods and Man . The New Naturalist: A Survey of British Natural History (3rd ed.). London: Collins. ISBN   978-0-00-213230-5. OCLC   1835850. OL   38560280M.
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  29. ↑ Wang, S.-R.; Zhang, H.; Chen, Y.-Z.; Zhang, Y.-D.; Li, D.-B.; Huang, Y.; Zhang, G.; Yang, J. (2022). "First Report of Black Spot Needle Blight of Pinus sylvestris var. mongolica Litv. Caused by Heterotruncatella spartii in China". Plant Disease. 106 (8): 2256. doi: 10.1094/PDIS-12-21-2667-PDN . PMID   35108070. S2CID   246487193.
  30. ↑ "Vegetable flannel". Webster's 1913 Dictionary. Retrieved 18 August 2014.
  31. ↑ Cole, G. S. (1892). A Complete Dictionary of Dry Goods and History of Silk, Cotton, Linen, Wool, and other Fibrous Substances, etc etc (revised ed.). Chicago: W. B. Conkey Company. pp. 365–366. OCLC   401856. OL   7231038M .
  32. ↑ Kershner, B.; Mathews, D.; Nelson, G.; Spellenberg, R.; Purinton, T.; Block, A.; Moore, G.; Thieret, J. W. (2008). National Wildlife Federation Field Guide to Trees of North America . New York: Sterling Publishing Company. p. 67. ISBN   978-1-4027-3875-3. OCLC   705626973. OL   11637505M.
  33. ↑ "Invasive Ontario Plants". Protect Our Water and Environmental Resources. Archived from the original on 26 April 2012. Retrieved 17 December 2011.
  34. ↑ "Species found in Michigan". Early Detection and Distribution Mapping System. Center for Invasive Species and Ecosystem Health at the University of Georgia . Retrieved 17 December 2011.
  35. ↑ Gleason, M.; Linit, M.; Zriba, N.; Donald, P.; Tisserat, N.; Giesler, L. (2000). Edwards, E. (ed.). Pine Wilt: A Fatal Disease of Exotic Pines in the Midwest (PDF). Sustainable Urban Landscapes. Iowa State University. SUL 9. Archived from the original on 15 May 2012. Retrieved 20 May 2023.{{cite book}}: CS1 maint: bot: original URL status unknown (link)
  36. ↑ "RHS Plantfinder - Pinus sylvestris (Aurea Group) 'Aurea'" . Retrieved 30 April 2018.
  37. ↑ "RHS Plant Selector – Pinus sylvestris 'Beuvronensis'" . Retrieved 6 February 2021.
  38. ↑ "RHS Plantfinder - Pinus sylvestris 'Frensham'" . Retrieved 30 April 2018.
  39. ↑ "RHS Plantfinder - Pinus sylvestris 'Gold Coin'" . Retrieved 25 April 2018.
  40. ↑ "AGM Plants - Ornamental" (PDF). Royal Horticultural Society. July 2017. p. 78. Retrieved 30 April 2018.
  41. ↑ Rick Steves Scotland (second edition) By Rick Steves
  42. ↑ The Pliocene flora of Frankfurt am Main, Germany: taxonomy, palaeoenvironments and biogeographic affinities by Zlatko Kvaček, Vasilis Teodoridis & Thomas Denk – Palaeobiodiversity and Palaeoenvironments, pages 647–703 (2020) – Springer Link – https://doi.org/10.1007/s12549-019-00391-6
  43. ↑ Oberpliocän-Flora aus den Baugruben des Klärbeckens bei Niederrad und Schleuse bei Höchst a M. T. Geyler F. Kinkelin. (Frankfort, 1887).
  44. 1 2 Cervantes S, Kesälahti R, Kumpula TA, Mattila TM, Helanterä H, Pyhäjärvi T (August 2023). "Strong Purifying Selection in Haploid Tissue-Specific Genes of Scots Pine Supports the Masking Theory". Mol Biol Evol. 40 (8). doi:10.1093/molbev/msad183. PMC   10457172 . PMID   37565532.
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