Temporal range: Early Permian–Recent
|Cycas rumphii with old and new male cones.|
Bessey 1907: 321.
Cycads // are seed plants typically characterized by a stout and woody (ligneous) trunk with a crown of large, hard and stiff, evergreen leaves. They usually have pinnate leaves. The individual plants are either all male or all female (dioecious). Cycads vary in size from having trunks only a few centimeters to several meters tall. They typically grow very slowly and live very long, with some specimens known to be as much as 1,000 years old. Because of their superficial resemblance, they are sometimes confused with and mistaken for palms or ferns, but are only distantly related to either.
Cycads are found across much of the subtropical and tropical parts of the world. They are found in South and Central America (where the greatest diversity occurs), Mexico, the Antilles, southeastern United States, Australia, Melanesia, Micronesia, Japan, China, Southeast Asia, India, Sri Lanka, Madagascar, and southern and tropical Africa, where at least 65 species occur. Some can survive in harsh semidesert climates (xerophytic), others in wet rain forest conditions, and some in both. Some can grow in sand or even on rock, some in oxygen-poor, swampy, bog-like soils rich in organic material, and some in both. Some are able to grow in full sun, some in full shade, and some in both. Some are salt tolerant (halophytes).
Cycads belong to the biological division Cycadophyta. The three extant families of cycads are Cycadaceae, Stangeriaceae, and Zamiaceae. Though they are a minor component of the plant kingdom today, during the Jurassic period, they were extremely common. They have changed little since the Jurassic, compared to some major evolutionary changes in other plant divisions.
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. They have been reported to fix nitrogen in association with a cyanobacterium living in the roots. These blue-green algae 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.
Cycads have a cylindrical trunk which usually does not branch. 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 (the leaf stalk has leaflets emerging from it as "ribs"), or have edges (margins) so deeply cut (incised) so as to appear compound. Some species have leaves that are bipinnate, which means the leaflets each have their own subleaflets, growing in the same form on the leaflet as the leaflets grow on the stalk of the leaf (self-similar geometry).
|Traditional view||Modern view|
The probable former range of cycads can be inferred from their global distribution. For example, the family Stangeriaceae only contains three extant species in Africa and Australia. Diverse fossils of this family have been dated to 135 mya, indicating that diversity may have been much greater before the Jurassic and late Triassic mass extinction events. However, the cycad fossil record is generally poor and little can be deduced about the effects of each mass extinction event on their diversity.
Instead, correlations can be made between the number of extant gymnosperms and angiosperms. It is likely that cycad diversity was affected more by the great angiosperm radiation in the mid-Cretaceous than by extinctions. Very slow cambial growth was first used to define cycads, and because of this characteristic the group could not compete with the rapidly growing, relatively short-lived angiosperms, which now number over 250,000 species, compared to the 947 remaining gymnosperms. It is surprising that the cycads are still extant, having been faced with extreme competition and five major extinctions. The ability of cycads to survive in relatively dry environments where plant diversity is generally lower, may explain their long persistence and longevity.
The cycad fossil record dates to the early Permian, 280 million years ago (mya). There is controversy over older cycad fossils that date to the late Carboniferous period, 300–325 mya. One of the first colonizers of terrestrial habitats, this clade probably diversified extensively within its first few million years, although the extent to which it radiated is unknown because relatively few fossil specimens have been found. The regions to which cycads are restricted probably indicate their former distribution in the Pangea before the supercontinents Laurasia and Gondwana separated. Recent studies have indicated the common perception of existing cycad species as living fossils is largely misplaced, with only Bowenia dating to the Cretaceous or earlier. Although the cycad lineage itself is ancient, most extant species have evolved in the last 12 million years.
The family Stangeriaceae (named for Dr. William Stanger, 1811–1854), consisting of only three extant species, is thought to be of Gondwanan origin, as fossils have been found in Lower Cretaceous deposits in Argentina, dating to 70–135 mya. The family Zamiaceae is more diverse, with a fossil record extending from the middle Triassic to the Eocene (54–200 mya) in North and South America, Europe, Australia, and Antarctica, implying the family was present before the break-up of Pangea. The family Cycadaceae is thought to be an early offshoot from other cycads, with fossils from Eocene deposits (38–54 mya) in Japan, China, and North America, indicating this family originated in Laurasia. Cycas is the only genus in the family and contains 99 species, the most of any cycad genus. Molecular data have recently shown Cycas species in Australasia and the east coast of Africa are recent arrivals, suggesting adaptive radiation may have occurred. The current distribution of cycads may be due to radiations from a few ancestral types sequestered on Laurasia and Gondwana, or could be explained by genetic drift following the separation of already evolved genera. Both explanations account for the strict endemism across present continental lines.
Overall species diversity 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 tropics. However, the peak in the northern tropics is largely due to Cycas in Asia and Zamia in the New World, whereas the peak in the southern tropics 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 cycad genera, and is dependent on the remaining species in each genus that did not follow the extinction pattern of their ancestors. Cycas is the only genus that has a broad geographical range and can thus be used to infer that cycads tend to live in the upper and lower tropics. This is probably because these areas have a drier climate with relatively cool winters; while cycads require some rainfall, they appear to be partly xerophytic. Potted specimens are found and thrive in global locations such as Canada, Russia, Finland and Chile.
||This article includes a list of references, but its sources remain unclear because it has insufficient inline citations. (July 2009)|
- Bessey, C.E. (1907). "A synopsis of plant phyla". Nebraska Univ. Stud. 7: 275–373.
- Brongniart, A. (1843). Énumération des genres de plantes cultivées au Musée d'histoire naturelle de Paris.
- Holtcamp, W. (2012). "The emerging science of BMAA: do cyanobacteria contribute to neurodegenerative disease?". Environmental health perspective 120 (3). doi:10.1289/ehp.120-a110. PMC 3295368. PMID 22382274.
- (Hermsen et al. 2006).
- Nagalingum, N. S. et al. Science. doi:10.1126/science.1209926. Missing or empty
- Hopkins, DJ; KR Johnson (December 1997). "First Record of cycad leaves from the Eocene Republic flora". Washington Geology 25 (4): 37.
- Chamberlain, C.J. (1965)  The Living Cycads, 1st ed. reprinted, New York : Hafner, 172 p., LCCN 65-027079
- Chaw, S.-M.; Parkinson, C.L.; Cheng, Y.; Vincent, T.M.; Palmer, J.D. (2000). "Seed plant phylogeny inferred from all three plant genomes: Monophyly of extant gymnosperms and origin of Gnetales from conifer" (PDF). Proceedings of the National Academy of Sciences 97 (8): 4086–4091. doi:10.1073/pnas.97.8.4086.
- Chaw, S.-M.; Walters, T.W.; Chang, C.-C.; Hu, S.-H.; Chen, S.-H. (2005). "A phylogeny of cycads (Cycadales) inferred from chloroplast matK gene, trnK intron, and nuclear rDNA ITS region". Molecular Phylogenetics and Evolution 37 (1): 214–234. doi:10.1016/j.ympev.2005.01.006.
- Chaw, S.-M.; Zharkikh, A.; Sung, H.-M.; Luu, T.-C.; Li, W.-H. (1997). "Molecular phylogeny of extant gymnosperms and seed plant evolution: Analysis of nuclear 18s rRNA sequences". Molecular Biology and Evolution 14 (1): 56–68. doi:10.1093/oxfordjournals.molbev.a025702.
- Crepet, W.L. (2000). "Progress in understanding angiosperm history, success, and relationships: Darwin's abominably 'perplexing phenomenon'". Proceedings of the National Academy of Sciences 97 (24): 12939–12941. doi:10.1073/pnas.97.24.12939.
- De Luca, Paulo (1990) "A Historical Perspective on Cycads from Antiquity to the Present", In: Stevenson, D. (Ed.) The Biology, Structure, and Systematics of the Cycadales, Memoirs of the New York Botanical Garden, 57, p. 1-7, ISBN 0-89327-350-3
- Donaldson, J.S. (2003) "Chapter 3: Regional Overview: Africa", In: Donaldson, J.S. (ed.), Cycads. Status Survey and Conservation Action Plan, IUCN/SSC Cycad Specialist Group, Gland; Cambridge : UKIUCN, ISBN 2-8317-0699-8, p. 9-19
- Donaldson, J. (2004) "Saving ghosts? The implications of taxonomic uncertainty and shifting infrageneric concepts in the cycadales for red listing and conservation planning", In: Walters,T. & Osborne, R. (eds), Cycad Classification: Concepts and Recommendations, Wallingford, UK : CABI, ISBN 0-85199-741-4, p. 13-22.
- Donaldson, J., Hill, K.D., & Stevenson, D.W. (2003a) "Chapter 2: Cycads of the World: An Overview", In: Donaldson, J.S. (ed.), Cycads. Status Survey and Conservation Action Plan, IUCN/SSC Cycad Specialist Group, Gland; Cambridge : UKIUCN, ISBN 2-8317-0699-8, p. 3-8
- Donaldson, J.S., Dehgan, A.P., Vovides, A.P., & Tang, W. (2003b) "Chapter 7: Cycads in trade and sustainable use of cycad populations", In: Donaldson, J.S. (ed.), Cycads. Status Survey and Conservation Action Plan, IUCN/SSC Cycad Specialist Group, Gland; Cambridge : UKIUCN, ISBN 2-8317-0699-8, p. 39-47
- Golding, J.S.; Hurter, P.J.H. (2003). "A Red List account of Africa's cycads and implications of considering life-history and threats". Biodiversity and Conservation 12 (3): 507–528. doi:10.1023/A:1022472801638.
- Gonzàlez-Astorga, J.; Vovides, A.P.; Ferrer, M.M.; Iglesias, C. (2003a). "Population genetics of Dioon edule Lindl. (Zamiaceae, Cycadales): biogeographical and evolutionary implications". Botanical Journal of the Linnean Society 80 (3): 457–467. doi:10.1046/j.1095-8312.2003.00257.x.
- Gonzàlez-Astorga, J.; Vovides, A.P.; Iglesias, C. (2003b). "Morphological and geographic variation in Dioon edule Lindl. (Zamiaceae): ecological and evolutionary implications". Botanical Journal of the Linnean Society 141 (4): 465–470. doi:10.1046/j.1095-8339.2003.00155.x.
- Gregory, T.J. & Chemnick, J. (2004) Hypotheses of the relationship between biogeography and speciation in Dioon (Zamiaceae). In: Walters,T. & Osborne, R. (eds), Cycad Classification: Concepts and Recommendations, Wallingford, UK : CABI, ISBN 0-85199-741-4, p. 137-148.
- Hermsen, E.; Taylor, E.; Taylor, T.; Stevenson, D. (2006). "Cataphylls of the middle Triassic cycad Antarcticycas schopfii and new insights into cycad evolution". American Journal of Botany 93 (724–738): 2006.
- Hill, C.R. (1990). "Ultrastructure of in situ fossil cycad pollen from the English Jurassic, with a description of the male cone Androstrobus balmei sp. nov". Review of Palaeobotany and Palynology 65 (1-4): 165–193. doi:10.1016/0034-6667(90)90067-S.
- Hill, K.D. (2003) "Chapter 4: Regional Overview: Australia", In: Donaldson, J.S. (ed.), Cycads. Status Survey and Conservation Action Plan, IUCN/SSC Cycad Specialist Group, Gland; Cambridge : UKIUCN, ISBN 2-8317-0699-8, p. 20-24
- Hill, K.D. (2004) "Character evolution, species recognition and classification concepts in the cycadaceae", In: Walters,T. & Osborne, R. (eds), Cycad Classification: Concepts and Recommendations, Wallingford, UK : CABI, ISBN 0-85199-741-4, p. 23-44. CABI, Oxford.
- Hill, K. & Stevenson, D. 1998–present. The Cycad Pages. http://plantnet.rbgsyd.nsw.gov.au/PlantNet/cycad/
- Hill, K.D.; Stevenson, D.W.; Obsorne, R. (2004). "The World List of Cycads". Botanical Review 70 (2): 274–298. doi:10.1663/0006-8101(2004)070[0274:TWLOC]2.0.CO;2.
- Hopkins, DJ, Johnson, KR "First record of cycad leaves from the Eocene Republic flora." Washington Geology Vol. 25 No. 4 p. 37 http://www.nwpaleo.org/Resources/WA_Geology/WA_Geol_1997-CycadLeaves.html
- Jones, David L. (1993) Cycads of the World, Washington, D.C. : Smithsonian Institution Press, ISBN 1-56098-220-9. 2nd ed. republished (2002) as: Cycads of the World: ancient plants in today's landscape, Sydney : Reed New Holland, ISBN 1-876334-69-X
- Keppel, G.; Lee, S.W.; Hodgskiss, P.D. (2002). "Evidence for Long Isolation Among Populations of a Pacific Cycad: Genetic Diversity and Differentiation in Cycas seemannii A.Br. (Cycadaceae)". Journal of Heredity 93 (2): 133–139. doi:10.1093/jhered/93.2.133.
- Norstog, K.J. & Nicholls, T.J. (1998) Biology of Cycads, Ithaca : Cornell University Press, ISBN 0-8014-3033-X
- Stevenson, D (1992). "A formal classification of the extant cycads". Brittonia 44: 220–223. doi:10.2307/2806837.
- Stevenson, D., K. Norstog, & P. Fawcett. 1998. Pollination biology of cycads. In: Reproductive Biology: In systematics, conservation, and economic botany. Eds. S. Owens & P. Rudall. pp. 277–294. Royal Botanic Gardens, Kew.
- Walters, T., Osborne, R., & Decker, D. (2004) "We hold these truths…[dead link]", In: Walters,T. & Osborne, R. (eds), Cycad Classification: Concepts and Recommendations, Wallingford, UK : CABI, ISBN 0-85199-741-4, p. 1-11
- Whitelock, L.M. (2002) The Cycads, Portland, OR : Timber Press, ISBN 0-88192-522-5
|Wikimedia Commons has media related to Cycadophyta.|
- Palm Trees, Small Palms, Cycads, Bromeliads and tropical plants site with thousands of large, high quality photos of cycads and associated flora. Includes information on habitat and cultivation.
- Hill KD (1998–2004) The Cycad Pages, Royal Botanic Gardens Sydney. http://plantnet.rbgsyd.nsw.gov.au/PlantNet/cycad/index.html
- Gymnosperm Database: Cycads[dead link]
- Fairchild Tropical Botanic Garden- one of the largest collection of cycads in the world in Florida, U.S.A.
- Palm and Cycad Societies of Australia (PACSOA)
- The Cycad Society of South Africa[dead link]
- Cycad nitrogen fixation
- Cycad toxicity
- Cycads - Foto
- The Cult of the Cycads, New York Times Magazine article on cycad collectorship and cycad smuggling
- Cycads An annotated link directory