Lithocarpus

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Lithocarpus
Temporal range: Eocene–Recent
Lithocarpus edulis Nakai leaf acorn.jpg
Lithocarpus edulis, Kantō region, Japan
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Fagales
Family: Fagaceae
Subfamily: Quercoideae
Genus: Lithocarpus
Blume
Species

See text

Diversity
c. 340 species
Synonyms

PasaniaOerst.

Lithocarpus sp. - MHNT Lithocarpus sp. MHNT.BOT.2013.22.30.jpg
Lithocarpus sp. - MHNT
Lithocarpus sp. - MHNT Lithocarpus sp. MHNT.BOT.2013.22.33.jpg
Lithocarpus sp. - MHNT

Lithocarpus is a genus in the beech family, Fagaceae. Trees in this genus are commonly known as the stone oaks and differ from Quercus primarily because they produce insect-pollinated flowers on erect spikes and the female flowers have short styles with punctate stigmas. At current, around 340 species have been described, [1] mostly restricted to Southeast Asia. Fossils show that Lithocarpus formerly had a wider distribution, being found in North America and Europe during the Eocene to Miocene epochs. [2] The species extend from the foothills of the Hengduan Mountains, where they form dominant stands of trees, through Indochina and the Malayan Archipelago, crossing Wallace's Line and reaching Papua. In general, these trees are most dominant in the uplands (more than 1,000 m or 3,300 ft above sea level) and have many ecological similarities to the Dipterocarpaceae, the dominant lowland tree group. [3] These trees are intolerant of seasonal droughts, not being found on the Lesser Sunda Islands, despite their ability to cross numerous water barriers to reach Papua. [4] [5]

Contents

The North American tanoak or tanbark oak (Notholithocarpus densiflorus) was previously included in this genus but recent evidence indicates the similarities in flower and fruit morphology are due to convergent evolution. Both genetic and morphological evidence demonstrate that the tanoak is a distant relative to Asian stone oaks, and therefore tanoak has been moved into a new genus, Notholithocarpus . [6]

Lithocarpus trees are evergreen trees with leathery, alternate leaves, the margins of which are almost always entire, rarely toothed. The seed is a nut similar to an oak acorn with a cupule enclosing the basal part of the fruit. Cupules of stone oaks demonstrate a wide variety in the type and arrangement of lamellae and scales on the outside of the cupule, with some of them completely enclosing the nut, even becoming irregularly dehiscent in a few species. The seeds are often protected by a hard woody shell (hence the genus name, from Greek λίθος, lithos, "stone" and καρπός, karpos, "seed"). In some sections of the genus, the seed is embedded in the basal material of the fruit which becomes highly lignified and hard, lending greater mechanical protection to the seed, [7] creating a novel type of fruit. [8] The kernel is edible in some species (e.g. Lithocarpus edulis ), but inedible, and very bitter, in others.

Several of the species are very attractive ornamental trees, used in parks and large gardens in warm temperate and subtropical areas.

Classification

In 1948, Aimee Camus produced a comprehensive treatment of the two major genera in the family, [9] given the specimens available to her at the Natural History Museum in Paris. Because of the many collections available from the French colonies in subtropical and tropical Indochina, she worked extensively with stone oaks from the region. Most importantly, she provided the only existing infrageneric structure within the genus but unfortunately, many of the species from the Malesian region, south of the Isthmus of Kra, are not incorporated into this system.

Her classification system included 13 subgenera, including the subgenus Pasania which is by far the largest division within the genus. About 100 Asian species were treated separately in Pasania, at the genus level, and occasionally the old name persists on some herbarium sheets that have not been annotated. Several of the other subgenera possess fewer than ten species and have distinctive morphologies. Few of the Malesian species are treated in Camus' system and Soepadmo, who wrote the Flora Malesiana treatment, [10] [11] made no attempt to update or integrate these species into Camus' system, therefore a lot of work obviously remains to be done. Camus' system was highly detailed, as three levels of organization are recognized below the subgenus, but the classification is not systematic at the lowest level.

List of subgenera (No. of species in Camus' treatment): Castanicarpus (1); Corylopasania (2); Cryptostylis (1); Cyclobalanus (58); Cyrtobalanus (1); Eulithocarpus (11); Gymnobalanus (10); Liebmannia (3); Oerstedia (1); Pachybalanus (14); Pasania (209); Pseudosynaedrys (9); Synaedrys (15); indeterminate (12).

Early researchers into the family often suggested that the stone oaks were primitive in the family. An exhaustive study of the inflorescence and fruits of 73 species from eight of Camus' subgenera found that important development and evolutionary characters distinguish the major groups in the genus and indicate differences among the genera of the family.

Species

Lithocarpus pseudoreinwardtii Lithocarpus pseudoreinwardtii Blanco2.440-cropped.jpg
Lithocarpus pseudoreinwardtii
Lithocarpus hancei Lithocarpus hancei Ying Ke Ke 20200326195505 02.jpg
Lithocarpus hancei

Related Research Articles

<span class="mw-page-title-main">Oak</span> Tree or shrub in the genus Quercus

An oak is a hardwood tree or shrub in the genus Quercus of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisphere; it includes some 500 species, both deciduous and evergreen. Fossil oaks date back to the Middle Eocene. Molecular phylogeny shows that the genus is divided into Old World and New World clades, but many oak species hybridise freely, making the genus's history difficult to resolve.

<span class="mw-page-title-main">Fagaceae</span> Family of flowering plants

The Fagaceae are a family of flowering plants that includes beeches, chestnuts and oaks, and comprises eight genera with about 927 species. Fagaceae in temperate regions are mostly deciduous, whereas in the tropics, many species occur as evergreen trees and shrubs. They are characterized by alternate simple leaves with pinnate venation, unisexual flowers in the form of catkins, and fruit in the form of cup-like (cupule) nuts. Their leaves are often lobed, and both petioles and stipules are generally present. Their fruits lack endosperm and lie in a scaly or spiny husk that may or may not enclose the entire nut, which may consist of one to seven seeds. In the oaks, genus Quercus, the fruit is a non-valved nut called an acorn. The husk of the acorn in most oaks only forms a cup in which the nut sits. Other members of the family have fully enclosed nuts. Fagaceae is one of the most ecologically important woody plant families in the Northern Hemisphere, as oaks form the backbone of temperate forest in North America, Europe, and Asia, and are one of the most significant sources of wildlife food.

<i>Castanopsis</i> Genus of flowering plants

Castanopsis, commonly called chinquapin or chinkapin, is a genus of evergreen trees belonging to the beech family, Fagaceae. The genus contains about 140 species, which are today restricted to tropical and subtropical eastern Asia. A total of 58 species are native to China, with 30 endemic; the other species occur further south, through Indochina to Indonesia and the Philippines, mountainous areas of Taiwan, and also in Japan. The English name chinkapin is shared with other related plants, including the golden chinkapins of the Pacific United States, which are sometimes included within Castanopsis but are more often considered a separate but very closely related genus, Chrysolepis.

Lithocarpus hendersonianus is a tree species in the beech family Fagaceae. Trees in the genus Lithocarpus are commonly known as stone oaks and differ from Quercus primarily because they produce insect-pollinated flowers. This tree species is endemic to Peninsular Malaysia. It is categorized as Vulnerable by the IUCN Red List of Threatened Species, primarily due to habitat loss.

Quercus asymmetrica is an oak tree species in the beech family Fagaceae. It is found in China and northern Vietnam. It is placed in subgenus Cerris, section Cyclobalanopsis.

<i>Notholithocarpus</i> Species of flowering plant in the family Fagaceae

Notholithocarpus densiflorus, commonly known as the tanoak or tanbark-oak, is a broadleaf tree in the family Fagaceae, and the type species of the genus Notholithocarpus. It is native to the far western United States, particularly Oregon and California. It ranges from 15–40 meters in height, with a trunk diameter of 60–190 centimeters.

Quercus kouangsiensis is an uncommon Asian species of trees in the beech family Fagaceae. It has been found only in southern China, in the Provinces of Guangdong, Guangxi, Hunan, and Yunnan. It is placed in subgenus Cerris, section Cyclobalanopsis.

Lithocarpus caudatifolius is a tree in the beech family Fagaceae. The specific epithet caudatifolius means 'leaf with caudate apex'.

Lithocarpus coopertus is a tree in the family Fagaceae. The specific epithet coopertus means 'covered over', referring to the acorn.

Lithocarpus daphnoideus is a tree in the family Fagaceae. The specific epithet daphnoideus means "like Daphne", referring to the genus Daphne and its leaves.

Lithocarpus encleisacarpus is a tree in the beech family Fagaceae. The specific epithet is from the Greek meaning 'enclosed fruit', referring to the acorns and cupules. The cupule is not fused to the nut though and often becomes irregularly dehiscent. The degree to which the nut is enclosed by the cupule varies across its geographic range. Trees in Lithocarpus are commonly known as the stone oaks and differ from Quercus primarily because they produce insect-pollinated flowers.

Quercus langbianensis is an uncommon oak tree species in the family Fagaceae. It is placed in subgenus Cerris, section Cyclobalanopsis, the ring-cupped oaks. These differ from other Quercus groups in that they have acorns with distinctive cups: usually with substantial rings, made-up of scales that have grown together. This species can be found in sub-tropical and tropical seasonal forests of Vietnam.

Quercus petelotii is the accepted name of an endemic oak tree species in the beech family Fagaceae; there are no known sub-species. It is placed in subgenus Cerris, section Cyclobalanopsis.

<i>Quercus arbutifolia</i> Species of oak tree

Quercus arbutifolia, synonym Quercus obovatifolia, is a species of oak in the beech family Fagaceae, native to southeast China and Vietnam. It is placed in Quercus sect. Cyclobalanopsis, the ring-cupped oaks. There are no known subspecies.

Lithocarpus jacobsii is a tree in the beech family Fagaceae. It is named for the Dutch botanist Marius Jacobs. Trees in Lithocarpus are commonly known as the stone oaks and differ from Quercus primarily because they produce insect-pollinated flowers.

Lithocarpus kalkmanii is a tree in the beech family Fagaceae. This species is named for the Dutch botanist Cornelis Kalkman.

Lithocarpus revolutus is a tree in the beech family Fagaceae. The name is derived from the way in which the margins of the leaves are typically rolled in upon themselves (revolute). Trees in Lithocarpus are commonly known as the stone oaks and differ from Quercus primarily because they produce insect-pollinated flowers.

<i>Quercus <span style="font-style:normal;">subg.</span> Quercus</i> Subgenus of Oak trees

Quercus subgenus Quercus is one of the two subgenera into which the genus Quercus was divided in a 2017 classification. It contains about 190 species divided among five sections. It may be called the New World clade or the high-latitude clade; most species are native to the Americas, the others being found in Eurasia and northernmost North Africa.

Quercus donnaiensis is a critically endangered species of oak trees, endemic to central Vietnam. This species belongs the Asian sub-genus of Quercus within the family Fagaceae: which have acorns with distinctive cups, characterised by growing rings of scaly protrusions; it was considered a synonym of Quercus langbianensis, but now included by Binh, Ngoc et al. (2018) in the "Q. langbianensis complex".

References

  1. ↑ Strijk, J.S. (September 4, 2018). "Lithocarpus - On: asianfagaceae.com – The complete database for information on the evolutionary history, diversity, identification and conservation of over 700 Species of Asian trees". Asian Fagaceae. Retrieved May 4, 2021.
  2. ↑ Liu, Xiaoyan; Song, Hanzhang; Jin, Jianhua (2020-08-14). "Diversity of Fagaceae on Hainan Island of South China During the Middle Eocene: Implications for Phytogeography and Paleoecology". Frontiers in Ecology and Evolution. 8: 255. doi: 10.3389/fevo.2020.00255 . ISSN   2296-701X.
  3. ↑ Ashton, Peter S. (1988). "Dipterocarp biology as a window to the understanding of tropical forest structure". Annual Review of Ecology and Systematics. 19: 347–370. doi:10.1146/annurev.es.19.110188.002023.
  4. ↑ Cannon, Charles H.; Manos, Paul S. (2003). "Phylogeography of the Southeast Asian stone oaks (Lithocarpus)" (PDF). Journal of Biogeography. 30 (2): 211–226. Bibcode:2003JBiog..30..211C. doi:10.1046/j.1365-2699.2003.00829.x. S2CID   86444402 . Retrieved 2020-03-23.
  5. ↑ Strijk, Joeri S.; Binh, Hoàng Thi; Ngoc, Nguyen Van; Pereira, Joan T.; Slik, J. W. Ferry; Sukri, Rahayu S.; Suyama, Yoshihisa; Tagane, Shuichiro; Wieringa, Jan J.; Yahara, Tetsukazu; Hinsinger, Damien D. (2020-05-22). "Museomics for reconstructing historical floristic exchanges: Divergence of stone oaks across Wallacea". PLOS ONE. 15 (5): e0232936. Bibcode:2020PLoSO..1532936S. doi: 10.1371/journal.pone.0232936 . ISSN   1932-6203. PMC   7244142 . PMID   32442164.
  6. ↑ Manos, Paul S.; Cannon, Charles H.; Oh, Sang-Hun (2008). "Phylogenetic relationships and taxonomic status of the paleoendemic Fagaceae of Western North America: recognition of a new genus, Notholithocarpus" (PDF). Madroño. 55 (3): 181–190. doi:10.3120/0024-9637-55.3.181. S2CID   85671229. Archived from the original (PDF) on 2017-03-20. Retrieved 2014-03-05.
  7. ↑ Chen, Xi; Cannon, Charles H.; Conklin-Brittan, Nancy L. (2012). "Evidence for a Trade-Off Strategy in Stone Oak (Lithocarpus) Seeds between Physical and Chemical Defense Highlights Fiber as an Important Antifeedant". PLOS ONE. 7 (3): e32890. Bibcode:2012PLoSO...732890C. doi: 10.1371/journal.pone.0032890 . PMC   3310818 . PMID   22457722.
  8. ↑ Cannon, Charles H.; Manos, Paul S. (2000). "The Bornean "Lithocarpus" Bl. section Synaedrys (Lindl.) Barnett (Fagaceae): its circumscription and description of a new species" (PDF). Botanical Journal of the Linnean Society. 133 (3): 343–357. doi:10.1006/bojl.1999.0325 . Retrieved 2020-03-20.
  9. ↑ Camus, Aimee (1948). Les chenes. Monographie des genres Quercus et Lithocarpus, Atlas vol. 3. Encyclopedie Économique de Sylviculture. Vol. 7. pp. 152–165. OCLC   496212700.
  10. ↑ Soepadmo, E. (1970). Florae Malesianae precursores XLIX. Malesian species of Lithocarpus BL. (Fagaceae). Reinwardtia. Vol. 8. pp. 197–308. OCLC   475354541.
  11. ↑ Soepadmo, E. (1972). "Fagaceae". Flora Malesiana: Series I - Spermatophytes. 7: 265–403. OCLC   8642966464.
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