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Trochodendraceae

From Wikipedia, the free encyclopedia
Family of flowering plants

Trochodendraceae
Temporal range:Paleocene–Recent
Trochodendron aralioides
Scientific classificationEdit this classification
Kingdom:Plantae
Clade:Tracheophytes
Clade:Angiosperms
Clade:Eudicots
Order:Trochodendrales
Takhtajan exCronquist
Family:Trochodendraceae
Eichler
Genera

Trochodendraceae is the onlyfamily offlowering plants in the orderTrochodendrales. It comprises two extantgenera, each with a singlespecies[1] along with up to five additional extinct genera and a number of extinct species.[2] The living species are native to south east Asia. The two living species (Tetracentron sinense andTrochodendron aralioides) both have secondary xylem withoutvessel elements, which is quite rare in angiosperms. As the vessel-free wood suggests primitiveness, these two species have attracted much taxonomic attention.

Description

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Tetracentron andTrochodendron are deciduous or evergreentrees, which grow to between 20–30 m (66–98 ft) tall, withTrochodendron sometimes sporting umbrella-shaped branches.

  • Leaves in spirals at the end of the branches (umbrella-like appearance,Trochodendron) or separate (Tetracentron), simple, serrulate or crenulate, with chloranthoid teeth, palmately or pinnately divided, brochidodromous or actinodromous, ovate orobovate, with a cordate to cuneate base and acuminate apex, stalked, with thinstipules fused with the petiole (Tetracentron) or absent (Trochodendron). Idioblasts present, large, branched, sclerenchymatous inTrochodendron and secretory inTetracentron.Stomata laterocytic or cyclocytic,hypostomatic.
  • Stems without xylematic vessels, with tracheids, xylem rays heterogeneous, uni- and multi-seriate, branches clearly differentiated in unifoliate brachyblasts and macroblasts with distichousphyllotaxis (Tetracentron), withnodes (1-)3(-multi)-lacunar, with (1)3(−7) leaf stems.
  • Hermaphroditic or androdioeciousplants.
  • TerminalInflorescence in erect, aggregated racemiforms (botryoid or small panicles) (Trochodendron) or defined, axillary, multi-floral amentoid spikes with the flower in whorls of 4 (Tetracentron).Bracts and bracteoles present or absent.
  • Perfectflowers, actinomorphic or dissymmetric, yellowish. Short, sub-conical, or hollowreceptacle. Hypogynous disk absent. Reduced, very thinperianth, of 4tepals in 2 decussate whorls (Tetracentron), or at most in a recognizable preantheric state (Trochodendron).Androecium of 4 decussatestamens in pairs of 40–70 in a spiral, non-versatile, basifixed, tetrasporangial, latrorso, apiculateanthers,dehiscence along 2 longitudinal valves in the theca.Gynoecium superior (Tetracentron) to slightly semi-inferior (Trochodendron), of 4–11(−17)carpels, syncarpous (alternating with the stamens inTetracentron) to semicarpous, the dorsal part of the ovary expanded horizontally in the anthesis, abaxially nectariferous, with sunken stomata, freestyles (stylodious), dry, papillose, decurrent ventralstigmas, 5–30 anatropous, apotropous, bitegmicous, crassinucelate, pendulousovules per carpel,placentation marginal in 2 series or apicoaxial.
  • Fruit in ventricidal or slightly loculicidalcapsule or an aggregate of dorsally and ventrally dehiscent semicarpicalfollicles, with basal and external styles.
  • Small, flattened, taperedseeds, 3–4 mm in length, with lateral, apical, chalazal wings, with thintesta, with abundant, oily and proteinaceousendosperm, smallembryo, with 2 cotyledons.
  • Pollen in small, granular, spheroidal, tricolpate, tectated-columelliform monads (10–20 μm in diameter), the surface with interwoven bars parallel to the edges of the colpus, which are granular.
  • Chromosomal number:2n = 48 inTetracentron and2n = 38, 40 inTrochodendron.

Ecology

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Pollination is probably myriophyllous, even thoughTetracentron shows a clear anemophilous syndrome. The pulverulent seeds are dispersed by the wind (anemochory).Trochodendron is present in both protandrous and protogynous forms that are self-compatible.

The plants are found in wooded formations,Trochodendron between 300 m and 2.700 m above sea level andTetracentron between 1.100 m and 3.600 m above sea level.

Phytochemistry

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Flavonoids (quercetin and kaempferol) andproanthocyanidins (cyanidin anddelphinidin) are present.Epicuticular waxes are basically composed of nonacosan-10-ol.[citation needed]Tetracentron containschalcones ordihydrochalcones.Trochodendron containsmyricetin.Ellagic acid is absent.

Fossils

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Trochodendron and the fossil genusEotrochion are known from thePaleocene of Wyoming, which constitute the oldest records of the family.[2]

A diverse assemblage of trochodendraceous species have been described from theEocene Okanagan Highlands in British Columbia and Northeastern Washington. The monotypic generaParaconcavistylon andPentacentron are accompanied byTetracentron hopkinsii and theTrochodendroninfructescence morphospeciesTr. drachukii plus the foliage morphospeciesTr. nastae.[3][2]

The fruit and wood genusNordenskioeldia, along with the associated leaf morphogenusZizyphoides have been considered part of Trochodendraceae, though phylogenetic analysis by Manchesteret al (2020) suggested they might be better placed as a basal stem lineage in the Trochodendrales, rather than as a crown group member of the family Trochodendraceae.[2]

Systematic position

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The Trocodendraceae are a group of flowering plants that include the cladeEudicotyledoneae. In previous systems they have been related with theCercidiphyllaceae and theEupteleaceae, with which they share some characteristics that can be considered symplesiomorphic or convergent and that have been excluded from the order Trochodendrales because of molecular data leaving the Trocodendraceae isolated. Based on molecular and morphological data, the APW (Angiosperm Phylogeny Website) considers that it constitutes the only family in the Order Trochodendrales (cf.AP-website).

Classification

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The current classification of Trochodendraceae is theAPG IV system published in 2016, which maintains the circumscription of Trochodendraceae used in theAPG III system published in October 2009. Unlike theAPG andAPG II systems, the later systems place the family as the only family in the order Trochodendrales. They also includesTetracentron, synonymizing Tetracentraceae fully with Trochodenraceae.[4][5]

TheAPG II system, of 2003 retained the classification used in the 1998APG system recognizing Trochodendraceae as a family. APG and APG II did not place the family in anorder, leaving it among the basal lineages of theeudicots. Both APG systems accepts this as a family of two modern species, but it does allow the option of separating out the familyTetracentraceae.

This segregation would lead to two families with one species each: Tetracentraceae withTetracentron sinense and Trochodendraceae withTrochodendron aralioides.

TheCronquist system, of 1981, accepted both Trochodendraceae and Tetracentraceae as families and placed these in the order Trochodendrales, in subclassHamamelidae, in classMagnoliopsida.

Taxa included

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The family includes two living genera with very different morphological characteristics:

  • Palmate leaves, with stipules, deciduous. Perianth of 4 tepals. Stamens 4. Carpels 4. Ovules 5–6 per carpel. Axillary inflorescence in amentoid spike.
TetracentronOliv., 1889. North-east India, Nepal, Bhutan, Burma, western and central China, Vietnam.
  • Pinnate leaves, without stipules, evergreen. Perianth absent. Stamens 40–70. Carpels (4-)6–11(−17). Ovules 15–30 per carpel. Terminal racemiform inflorescence, erect.
TrochodendronSiebold &Zucc., 1839. Japan, Taiwan, Korea.

References

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  1. ^Christenhusz, M. J. M. & Byng, J. W. (2016)."The number of known plants species in the world and its annual increase".Phytotaxa.261 (3):201–217.Bibcode:2016Phytx.261..201C.doi:10.11646/phytotaxa.261.3.1.
  2. ^abcdManchester, S. R.; Kvaček, Z.; Judd, W. S. (2020)."Morphology, anatomy, phylogenetics and distribution of fossil and extant Trochodendraceae in the Northern Hemisphere".Botanical Journal of the Linnean Society.195 (3):467–484.doi:10.1093/botlinnean/boaa046.
  3. ^Manchester, S.; Pigg, K. B.; Kvaček, Z; DeVore, M. L.; Dillhoff, R. M. (2018)."Newly recognized diversity in Trochodendraceae from the Eocene of western North America".International Journal of Plant Sciences.179 (8):663–676.Bibcode:2018IJPlS.179..663M.doi:10.1086/699282.S2CID 92201595.
  4. ^Angiosperm Phylogeny Group (2009).An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III.Botanical Journal of the Linnean Society161: 105–121.
  5. ^Angiosperm Phylogeny Group (2016)."An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV".Botanical Journal of the Linnean Society.181 (1):1–20.doi:10.1111/boj.12385.ISSN 0024-4074.

Further reading

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External links

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