Cycad

For the insect, see Cicada.

Cycadales Temporal range: Early Permian–Holocene PreꞒ Ꞓ O S D C P T J K Pg N
Cycas rumphii with old and new male strobili.
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Spermatophytes
Clade:	Gymnospermae
Division:	Cycadophyta Bessey 1907: 321.[2]
Class:	Cycadopsida Brongn.[1]
Order:	Cycadales Pers. ex Bercht. & J. Presl
Extant groupings
Cycadaceae Zamiaceae
Synonyms
Cycadofilicales Němejc 1950 Dioales Doweld 2001 Stangeriales Doweld 2001 Zamiales Burnett 1835

Cycads in South Africa

Cycads /ˈsaɪkædz/ are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or female. Cycads vary in size from having trunks only a few centimeters to several meters tall. They typically grow slowly^[3] and have long lifespans. Because of their superficial resemblance to palms or ferns, they are sometimes mistaken for them, but they are not closely related to either group. Cycads are gymnosperms (naked-seeded), meaning their unfertilized seeds are open to the air to be directly fertilized by pollination, as contrasted with angiosperms, which have enclosed seeds with more complex fertilization arrangements. Cycads have very specialized pollinators, usually a specific species of beetle. Both male and female cycads bear cones (strobili), somewhat similar to conifer cones.

Cycads have been reported to fix nitrogen in association with various cyanobacteria living in the roots (the "coralloid" roots).^[4] These photosynthetic bacteria produce a neurotoxin called BMAA that is found in the seeds of cycads. This neurotoxin may enter a human food chain as the cycad seeds may be eaten directly as a source of flour by humans or by wild or feral animals such as bats, and humans may eat these animals. It is hypothesized that this is a source of some neurological diseases in humans.^[5][6] Another defence mechanism against herbivores is the accumulation of toxins in seeds and vegetative tissues; through horizontal gene transfer, cycads have acquired a family of genes (fitD) from a microbial organism, most likely a fungus, which gives them the ability to produce an insecticidal toxin.^[7]

Cycads all over the world are in decline, with four species on the brink of extinction and seven species having fewer than 100 plants left in the wild.^{[8][better source needed]}

Description

Cycads have a rosette of pinnate leaves around a cylindrical trunk

Cycads have a cylindrical trunk which usually does not branch. However, some types of cycads, such as Cycas zeylanica, can branch their trunks. The apex of the stem is protected by modified leaves called cataphylls.^[9] Leaves grow directly from the trunk, and typically fall when older, leaving a crown of leaves at the top. The leaves grow in a rosette form, with new foliage emerging from the top and center of the crown. The trunk may be buried, so the leaves appear to be emerging from the ground, so the plant appears to be a basal rosette. The leaves are generally large in proportion to the trunk size, and sometimes even larger than the trunk.

The leaves are pinnate (in the form of bird feathers, pinnae), with a central leaf stalk from which parallel "ribs" emerge from each side of the stalk, perpendicular to it. The leaves are typically either compound (with leaflets emerging from the leaf stalk as "ribs"), or have edges (margins) so deeply cut (incised) so as to appear compound. The Australian genus Bowenia and some Asian species of Cycas, like Cycas multipinnata, C. micholitzii and C. debaoensis, have leaves that are bipinnate, the leaflets each having their own subleaflets, growing in the same form on the leaflet as the leaflets do on the stalk.^[10][11]

Confusion with palms

Due to superficial similarities in foliage and plant structure, cycads and palms are often mistaken for each other. They also can occur in similar climates. However, they belong to different phyla and as such are not closely related. The similar structure is the product of convergent evolution.

Beyond those superficial resemblances, there are a number of differences between cycads and palms. For one, both male and female cycads are gymnosperms and bear cones (strobili), while palms are angiosperms and so flower and bear fruit. The mature foliage looks very similar between both groups, but the young emerging leaves of a cycad resemble a fiddlehead fern before they unfold and take their place in the rosette, while the leaves of palms are just small versions of the mature frond. Another difference is in the stem. Both plants leave some scars on the stem below the rosette where there used to be leaves, but the scars of a cycad are helically arranged and small, while the scars of palms are a circle that wraps around the whole stem. The stems of cycads are also in general rougher and shorter than those of palms.^[12]

Taxonomy

The two extant families of cycads all belong to the order Cycadales, and are the Cycadaceae and Zamiaceae (including Stangeriaceae). These cycads have changed little since the Jurassic in comparison to some other plant divisions. Five additional families belonging to the Medullosales became extinct by the end of the Paleozoic Era.

Based on genetic studies, cycads are thought to be more closely related to Ginkgo than to other living gymnosperms. Both are thought to have diverged from each other during the early Carboniferous.^[13][14]

External phylogeny[13][14]	Internal phylogeny[15][16]
Cycads Ginkgo Conifers Anthophytes Bennettitales Gnetales Angiosperms Gymnosperms Cycads Ginkgo Conifers Gnetophytes Angiosperms (flowering plants) Traditional view Modern view	Cycads Cycadineae Cycadaceae Cycas Zamiineae Zamiaceae Diooideae Dioon Zamioideae Encephalarteae Macrozamia Lepidozamia Encephalartos Zamieae Bowenia Ceratozamia Stangeria Zamia Microcycas

Classification of the Cycadophyta to the rank of family.

• Class Cycadopsida Brongniart 1843
  • Order Cycadales Persoon ex von Berchtold & Presl 1820
    • Suborder Cycadineae Stevenson 1992
      • Family Cycadaceae Persoon 1807
        • Genus Cycas
    • Suborder Zamiineae Stevenson 1992
      • Family Zamiaceae Horaninow 1834
        • subfamily Diooideae Pilg. 1926
          • Tribe Diooeae Schuster
            • Genus Dioon
        • subfamily Zamioideae Stevenson 1992
          • Tribe Encephalarteae Miquel 1861
            • Genus Macrozamia
            • Genus Lepidozamia
            • Genus Encephalartos
          • Tribe Zamieae Miquel 1861
            • Genus Bowenia
            • Genus Ceratozamia
            • Genus Stangeria
            • Genus Zamia
            • Genus Microcycas

Fossil genera

The following extinct cycad genera are known:^[17]

• Amuriella Late Jurassic, Russian Far East (leaf fragments)
• Androstrobus Triassic to Cretaceous, worldwide (leaf form genus)
• Antarcticycas Middle Triassic, Antarctica (known from the whole plant)^[18]
• ?Anthrophyopsis Late Triassic, worldwide (leaf form genus, possibly a pteridospermatophyte)^[19]
• Apoldia Triassic-Jurassic, Europe
• Archaeocycas Early Permian, Texas (leaf with sporophylls)
• Aricycas Late Triassic, Arizona (leaf form genus)
• Beania (=Sphaereda), Triassic to Jurassic, Europe & Central Asia (leaf form genus)
• Behuninia Late Jurassic, Colorado & Utah (fruiting structures)
• Bucklandia Middle Jurassic to Early Cretaceous, Europe and India (leaf form genus)
• Bureja Late Jurassic, Russia
• Cavamonocolpites Early Cretaceous, Brazil (pollen)
• Crossozamia Early to Late Permian, China (leaf form genus)
• Ctenis Mesozoic-Paleogene, Worldwide (leaf form genus)
• Ctenozamites Triassic-Cretaceous, worldwide (leaf form genus)
• Cycadenia Triassic, Pennsylvania (trunks)
• Cycadinorachis Late Jurassic, India (rachis)
• Fascisvarioxylon Late Jurassic, India (petrified wood)
• Gymnovulites, Latest Cretaceous/earliest Paleocene, India (seed)
• Heilungia, Late Jurassic to early Cretaceous, Russia & Alaska (leaf form genus)
• Leptocycas Late Triassic, North Carolina & China (known from the whole plant)^[20]
• Mesosingeria, Jurassic to Early Cretaceous, Antarctica & Argentina (leaf form genus)
• Michelilloa, Late Triassic, Argentina (stem)
• ?Nikania, Early Cretaceous, Russia (leaf fragments)
• ?Nilssonia, Middle Permian to Late Cretaceous, worldwide (leaf form genus) (possibly not a cycad)^[21]
• ?Nilssoniocladus, Early to Late Cretaceous, United States & Russia (stems, likely associated with Nilssonia, possibly deciduous)^[22]
• Palaeozamia, Middle Jurassic, England
• Paracycas, Middle Jurassic to Late Jurassic, Europe and Central Asia
• ?Phasmatocycas, Late Carboniferous to Early Permian, Kansas, Texas & New Mexico (leaf with sporophylls)^[23]
• Pleiotrichium, Late Cretaceous, Germany (leaf)
• Pseudoctenis, Late Permian to Late Cretaceous, worldwide (leaf form genus)
• Sarmatiella, Late Triassic, Ukraine
• Stangerites, Late Triassic to Early Jurassic, Virginia and Mexico (leaf form genus)
• Sueria, Early Cretaceous, Argentina (leaf)
• Taeniopteris, Carboniferous to Cretaceous, worldwide (polyphyletic leaf form genus, also includes bennettitales and marattialean ferns)

Fossil record

Bowenia spectabilis : plant with single frond in the Daintree rainforest, north-east Queensland

Leaves and strobilus of Encephalartos sclavoi

The oldest probable cycad foliage is known from the latest Carboniferous-Early Permian of South Korea and China, such as Crossozamia. Unambiguous fossils of cycads are known from the Early-Middle Permian onwards.^[24] Cycads were generally uncommon during the Permian.^[25] The two living cycad families are thought to have split from each other sometime between the Jurassic^[15] and Carboniferous.^[26] Cycads are thought to have reached their apex of diversity during the Mesozoic.^[27] Although the Mesozoic is sometimes called the "Age of Cycads," some other groups of extinct seed plants with similar foliage, such as Bennettitales and Nilssoniales, that are not closely related, may have been more abundant.^[28] The oldest records of the modern genus Cycas are from the Paleogene of East Asia.^[29] Fossils assignable to Zamiaceae are known from the Cretaceous,^[28] with fossils assignable to living genera of the family known from the Cenozoic.^[16]

Petrified cycad fossil, New York Botanical Garden

Distribution

See also: List of cycad species by country

The living cycads are found across much of the subtropical and tropical parts of the world, with a few in temperate regions such as in Australia.^[30] The greatest diversity occurs in South and Central America.^{[citation needed]} They are also found in Mexico, the Antilles, southeastern United States, Australia, Melanesia, Micronesia, Japan, China, Southeast Asia, Bangladesh, India, Sri Lanka, Madagascar, and southern and tropical Africa, where at least 65 species occur. Some can survive in harsh desert or semi-desert climates (xerophytic),^[31] others in wet rain forest conditions,^[32] and some in both.^[33] Some can grow in sand or even on rock, some in oxygen-poor, swampy, bog-like soils rich in organic material.^{[citation needed]} Some are able to grow in full sun, some in full shade, and some in both.^{[citation needed]} Some are salt tolerant (halophytes).^{[citation needed]}

Species diversity of the extant cycads peaks at 17˚ 15"N and 28˚ 12"S, with a minor peak at the equator. There is therefore not a latitudinal diversity gradient towards the equator but towards the Tropic of Cancer and the Tropic of Capricorn. However, the peak near the northern tropic is largely due to Cycas in Asia and Zamia in the New World, whereas the peak near the southern tropic is due to Cycas again, and also to the diverse genus Encephalartos in southern and central Africa, and Macrozamia in Australia. Thus, the distribution pattern of cycad species with latitude appears to be an artifact of the geographical isolation of the remaining cycad genera and their species, and perhaps because they are partly xerophytic rather than simply tropical.

Cultural significance

Nuts of the Cycas orientis (nyathu) are coveted by the Yolngu in Australia's Arnhem Land as a source of food. They are harvested on their dry season to leach its poison under water overnight before ground into a paste, wrapped under bark and cooked on open fire until done.^[34]

In Vanuatu, the cycad is known as namele and is an important symbol of traditional culture. It serves as a powerful taboo sign,^[35] and a pair of namele leaves appears on the national flag and coat of arms. Together with the nanggaria plant, another symbol of Vanuatu culture, the namele also gives its name to Nagriamel, an indigenous political movement.

See also

• Fossil Cycad National Monument, formerly in the U.S. state of South Dakota

References

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10. Rutherford, Catherine. CITES and Cycads: A User's Guide (PDF). Royal Botanic Gardens, Kew.
11. Lariushin, Boris (19 January 2013). Cycadaceae Family. Lulu.com. ISBN 9781300654537.
12. Tudge, Colin (2006). The Tree. New York: Crown Publishers. pp. 70–72, 139–148. ISBN 978-1-4000-5036-9.
13. Wu, Chung-Shien; Chaw, Shu-Miaw; Huang, Ya-Yi (January 2013). "Chloroplast phylogenomics indicates that Ginkgo biloba is sister to cycads". Genome Biology and Evolution. 5 (1): 243–254. doi:10.1093/gbe/evt001. ISSN 1759-6653. PMC 3595029. PMID 23315384.
14. Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu (19 July 2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. doi:10.1038/s41477-021-00964-4. ISSN 2055-0278. PMID 34282286. S2CID 236141481.
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18. Hermsen, Elizabeth J.; Taylor, Edith L.; Taylor, Thomas N. (January 2009). "Morphology and ecology of the Antarcticycas plant". Review of Palaeobotany and Palynology. 153 (1–2): 108–123. Bibcode:2009RPaPa.153..108H. doi:10.1016/j.revpalbo.2008.07.005.
19. Xu, Yuanyuan; Popa, Mihai Emilian; Zhang, Tingshan; Lu, Ning; Zeng, Jianli; Zhang, Xiaoqing; Li, Liqin; Wang, Yongdong (1 September 2021). "Re-appraisal of Anthrophyopsis (Gymnospermae): New material from China and global fossil records". Review of Palaeobotany and Palynology. 292: 104475. Bibcode:2021RPaPa.29204475X. doi:10.1016/j.revpalbo.2021.104475. ISSN 0034-6667.
20. Zhang, Jian-Wei; Yao, Jian-Xin; Chen, Jia-Rui; Li, Cheng-Sen (25 May 2010). "A new species of Leptocycas (Zamiaceae) from the Upper Triassic sediments of Liaoning Province, China". Journal of Systematics and Evolution. 48 (4): 286–301. doi:10.1111/j.1759-6831.2010.00079.x.
21. Vajda, Vivi; Pucetaite, Milda; McLoughlin, Stephen; Engdahl, Anders; Heimdal, Jimmy; Uvdal, Per (August 2017). "Molecular signatures of fossil leaves provide unexpected new evidence for extinct plant relationships". Nature Ecology & Evolution. 1 (8): 1093–1099. Bibcode:2017NatEE...1.1093V. doi:10.1038/s41559-017-0224-5. ISSN 2397-334X. PMID 29046567. S2CID 3604369.
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24. Spiekermann, Rafael; Jasper, André; Siegloch, Anelise Marta; Guerra-Sommer, Margot; Uhl, Dieter (June 2021). "Not a lycopsid but a cycad-like plant: Iratinia australis gen. nov. et sp. nov. from the Irati Formation, Kungurian of the Paraná Basin, Brazil". Review of Palaeobotany and Palynology. 289: 104415. Bibcode:2021RPaPa.28904415S. doi:10.1016/j.revpalbo.2021.104415. S2CID 233860955.
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External links

• "Palm Trees, Small Palms, Cycads, Bromeliads and tropical plants". Archived from the original on 20 May 2014. Site with thousands of large, high quality photos of cycads and associated flora. Includes information on habitat and cultivation. (Site is dead.)
• "The Cycad Pages". Royal Botanic Gardens Sydney. Archived from the original on 29 March 2021.
• "Gymnosperm Database: Cycads". Archived from the original on 10 August 2010.
• "Fairchild Tropical Botanic Garden". Archived from the original on 23 February 2015. One of the largest collections of cycads in the world in Florida, U.S.A.
• "Main Page". Palm and Cycad Societies of Australia (PACSOA). Retrieved 18 June 2023.
• "Welcome". The Cycad Society of South Africa. Retrieved 18 June 2023.
• David Dalton (1997). "Part I. The range of organisms that can fix nitrogen". Nitrogen Fixation. Retrieved 18 June 2023. Cycad nitrogen fixation
• "Cycad toxicity". The Cycad Pages. Royal Botanic Gardens Sydney. Archived from the original on 17 July 2020.
• Lauren Kessler (28 August 2005). "The Cult of the Cycads". The New York Times Magazine. Archived from the original on 4 June 2022. Magazine article on cycad collectorship and cycad smuggling.