Zamia
| Zamia | |
|---|---|
| |
| Zamia furfuracea | |
Scientific classification 
| |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Gymnospermae |
| Division: | Cycadophyta |
| Class: | Cycadopsida |
| Order: | Cycadales |
| Family: | Zamiaceae |
| Subtribe: | Zamiinae |
| Genus: | Zamia L.[1] |
| Type species | |
| Zamia pumila | |
| Synonyms[2] | |
| |
Zamia is a genus of cycad of the family Zamiaceae, native to North America from the United States (in Georgia and Florida) throughout the West Indies, Central America, and South America as far south as Bolivia.[2][3][4][5] The genus is considered to be the most ecologically and morphologically diverse of the cycads, and is estimated to have originated about 68.3 million years ago.[6]
Description
[edit]The genus comprises deciduous shrubs with aerial or subterranean circular stems, often superficially resembling palms. They produce spirally arranged, pinnate leaves which are pubescent, at least when young, having branched and simple, transparent and coloured hairs. The articulated leaflets lack a midrib, and are broad with subparallel dichotomous venation. Lower leaflets are not reduced to spines, though the petioles often have prickles. The emerging leaves of many Zamia species are striking, some emerging with a reddish or bronze cast (Z. roezlii being an example). Zamia picta is even more distinctive, being the only truly variegated cycad (having whitish/yellow speckles on the leaves).[7]
Reproduction
[edit]Zamia sporophylls are born in vertical rows in cones, and the megasporophyll apices are faceted or flattened, not spinose. The fleshy seeds are subglobular to oblong or ellipsoidal, and are red, orange, yellow or rarely white. The endosperm is haploid, derived from the female gametophyte. The embryo is straight, with two cotyledons that are usually united at the tips and a very long, spirally twisted suspensor. The sperm of members from the genus are large, as is typical of cycads, and Z. roezlii is an example; its sperm are approximately 0.4 mm long and can be seen by the unaided eye.[8]
Preferred habitat
[edit]
All the species of Zamia produce leafy crowns of foliage that make them choice garden specimens and most varieties branch heavily in age to produce handsome clumps. With a few exceptions, most Zamia species are found in warm, humid, tropical rainforest habitats, growing in the forest understory. However, many species are still fairly adaptable, performing quite well in cultivation, especially in subtropical areas. All species need good drainage and protection from the cold.
Ecology
[edit]At least one species, Z. pseudoparasitica, grows as an epiphyte in the branches of trees.
Phylogeny
[edit]| Phylogeny of Zamia[9][10] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Geographic groups
[edit]Zonneveld and Lindström (2016) measured genome size in 71 species of Zamia and found support for three geographical groupings. Variation in genome size of Zamia species is fairly small compared to other many other genera, with the ratio of largest to smallest just 1.36, but the authors found significant differences in genome sizes between three geographical areas. Species in Mega Mexico, including the northern part of Central America, had the largest average genome size. Species in South America, plus Costa Rica and Panama, had the smallest average genome size, while species in the Caribbean Islands and Florida had an intermediate genome size.[11]
Calonje, et al. (2019) analyzed the DNA from 70 species of Zamia, finding support for five geographically distinct clades. A clade including the species found on the Caribbean islands and in Florida is sister to the rest of the genus. The Fischeri clade includes three species found in Mexico, and is sister to the rest of the genus. The Mesoamerica clade includes all species found in Mesoamerica (north of Costa Rica), except for the Fischeri clade and the single species Z. sosonuscensis. The Isthmus clade includes species found in Costa Rica and Panama. The species in South America form another clade, which is sister to the Isthmus clade.[12]
Lindstrom et al. (2024) analyzed transcriptomes from 77 species of Zamia finding support for seven clades of the genus occupying distinct geographical ranges. Clade I is a strongly monophyletic clade that includes all nine of the species of the Caribbean islands and Florida. Clade II (the Fischeri clade), consists of three species found in Veracruz, Hidalgo, Querétaro, San Luis Potosí, and Tamaulipas states in Mexico. This clade is a sister to Clades I and III. Clade III (Mega Mexico) is divided into the sub-clades III-A and III-B. Clade III-A includes 15 species found in Mexico and Central America. Clade III-B consists of seven species found in Honduras, Guatemala, and Belize, which are believed to have originated in southern Mexico. Clade IV consists of the single species Z. sosonuscensis found in cloud forests in Chiapas state in Mexico. Clade V (the Isthmus clade) includes 15 species found in Costa Rica and Panama. Clade VI includes 12 species found in southernmost Panama and west of the Andes in Colombia and Ecuador. Clade VII consists of four closely related species in northern Columbia (the Manicata clade) and 13 species east of the Andes in Venezuela, Colombia, Ecuador, Peru, Brazil, and Bolivia.[13]
Zamia pumila species complex
[edit]All of the species in Lindstrom et al.'s Clade I above are also in the Zamia pumila species complex. Controversy has long existed over the classification and circumscription of Zamia species in the southeastern United States and West Indies. At one extreme, Eckenwalder (1980) included all Zamia in the Caribbean in the single species Z. pumila, incorporating 27 previously described species (not all of which were valid or accepted) into the subspecies Z. pumila subsp. pumila, and five such species into the subspecies Z. pumila subsp. pygmaea.[14] Eckenwalder's classification is no longer generally accepted, and a monophyletic species complex consisting of nine species is now accepted, including Z. pumila, seven species corresponding to combinations of the species subsumed into Eckenwalder's Z. pumila subsp. pumila, and Z. pygmaea, consisting of the former species placed by Eckenwalder in Z. pumila subsp. pygmaea.[15][16]
Other species complexes
[edit]- Z. katzeriana, Mexico
- Z. loddigesii
- Z. poeppigiana, Colombia
- Z. skinneri, Central America
Species
[edit]Accepted species:[17]
| Species | Clade[13]: distribution | IUCN status |
|---|---|---|
| Zamia acuminata Oerst.ex Dyer | V (Isthmus): Costa Rica, Nicaragua, Panama | Vulnerable |
| Zamia amazonum D.W.Stev. | VII (East of Andes): Brazil, Colombia, Ecuador, Peru, Venezuela | Least concern |
| Zamia amplifolia Hort.Bull ex Mast. | VI (West of Andes): Colombia | Endangered |
| Zamia angustifolia Jacq. | I (Caribbean): Bahamas, Cuba | Vulnerable |
| Zamia boliviana (Brongn.) A.DC. | VII (East of Andes): Bolivia, Brazil (Mato Grosso) | Least concern |
| Zamia brasiliensis Calonje & Segalla[18] | (unknown): Brazil (Mato Grosso, Rondônia) | Not evaluated |
| Zamia chigua Seem. | VI (West of Andes): Colombia | Near threatened |
| Zamia cremnophila Vovides, Schutzman & Dehgan | III-A (Mega Mexico): Mexico (Tabasco) | Endangered |
| Zamia cunaria Dressler & D.W.Stev. | VI (West of Andes): Panama | Endangered |
| Zamia decumbens Calonje, Meerman, M.P. Griff. & Hoese | III-B (Mega Mexico) Belize | Endangered |
| Zamia disodon D.W.Stev. & Sabato | VII (Manicata): Colombia | Endangered |
| Zamia dressleri D.W.Stev. | V (Isthmus): Panama | Endangered |
| Zamia elegantissima Schutzman, Vovides & R.S.Adams | V (Isthmus): Panama | Endangered |
| Zamia encephalartoides D.W. Stev. | VII (East of Andes): Colombia | Endangered |
| Zamia erosa O.F.Cook & G.N.Collins | I (Caribbean): Cuba, Jamaica, Puerto Rico | Vulnerable |
| Zamia fairchildiana L.D.Gómez | V (Isthmus): Panama, Costa Rica | Near threatened |
| Zamia fischeri Miq. ex Lem. | II (Fischeri): Mexico (Hidalgo, San Luis Potosí, Querétaro, Tamaulipas, Veracruz) | Endangered |
| Zamia furfuracea L.f. cardboard palm | III-A (Mega Mexico): Mexico (Veracruz) | Endangered |
| Zamia gentryi Dodson | VI (West of Andes): Ecuador | Endangered |
| Zamia gomeziana R.H.Acuña | (unknown): Costa Rica | Near threatened |
| Zamia grijalvensis Pérez-Farr., Vovides & Mart.-Camilo | III-A (Mega Mexico): Mexico (Chiapas) | Critically endangered |
| Zamia hamannii A.S.Taylor, J.L.Haynes & Holzman | V (Isthmus): Panama | Vulnerable |
| Zamia herrerae Calderón & Standl. | III-A (Mega Mexico): Mexico (Chiapas), El Salvador, Guatemala, Honduras, Nicaragua | Vulnerable |
| Zamia huilensis Calonje, H.E.Esquivel & D.W.Stev. | VII (East of Andes): Colombia | Endangered |
| Zamia hymenophyllidia D.W.Stev. | VII (East of Andes): Colombia, Peru | Least concern |
| Zamia imbricata Calonje & Castro[19] | VII (Manicata): Columbia | Not evaluated |
| Zamia imperialis A.S.Taylor, J.L.Haynes & Holzman | V (Isthmus): Panama | Endangered |
| Zamia incognita A.Lindstr. & Idárraga | (unknown): Colombia | Endangered |
| Zamia inermis Vovides, J.D.Rees & Vázq.Torres | II (Fischeri): Mexico (Veracruz) | Critically endangered |
| Zamia integrifolia L.f. - coontie palm/Florida arrowroot | I (Caribbean): Bahamas, Cayman Islands, Cuba, Puerto Rico, United States (Florida, Georgia) | Near threatened |
| Zamia ipetiensis D.W.Stev. | VI (West of Andes): Panama | Endangered |
| Zamia katzeriana (Regel) E.Rettig | III-A (Mega Mexico): Mexico | Critically endangered |
| Zamia lacandona Schutzman & Vovides | III-A (Mega Mexico): Mexico (Chiapas) | Endangered |
| Zamia lecointei Ducke | VII (East of Andes): Brazil (Pará), Colombia, Peru, Venezuela | Least concern |
| Zamia lindenii Regel ex André | VI (West of Andes): Ecuador, Peru | Endangered |
| Zamia lindleyi Warsz. ex A.Dietr. | V (Isthmus): Panama | Endangered |
| Zamia lindosensis Stevenson, Cárdenas & Castaño[20] | (unknown): Colombia | Not evaluated |
| Zamia loddigesii Miq. | III-A (Mega Mexico): Belize, El Salvador, Honduras, Mexico (Campeche, Chiapas, Veracruz, Puebla, Oaxaca, Hidalgo, Tabasco, Tamaulipas, Quintana Roo, Yucatán) | Near threatened |
| Zamia lucayana Britton | I (Caribbean): Bahamas | Critically endangered |
| Zamia macrochiera D.W.Stev. | (unknown): Peru | Endangered |
| Zamia magnifica Pérez-Farr., Gutt-Ortega & Calonje[21] | (unknown): Mexico (Oaxaca) | Not evaluated |
| Zamia manicata Linden ex Regel | VII (Manicata): Colombia, Panama | Near threatened |
| Zamia meermanii Calonje | III-A (Mega Mexico): Belize | Endangered |
| Zamia melanorrhachis D.W.Stev. | (unknown): Colombia | Endangered |
| Zamia montana A.Braun | VI (West of Andes): Colombia | Critically endangered |
| Zamia monticola Chamb. | III-B (Mega Mexico): Guatemala | Critically endangered |
| Zamia multidentata Calonje, Segalla & R.S.Pimenta[22] | (unknown): Brazil (Acre) | Not evaluated |
| Zamia muricata Willd. | VII (East of Andes): Colombia, Venezuela | Least concern |
| Zamia nana A.Lindstr., Calonje, D.W.Stev. & A.S.Taylor Panama[23] | V (Isthmus): Panama | Endangered |
| Zamia nesophila A.S.Taylor, J.L.Haynes & Holzman | V (Isthmus): Panama | Critically endangered |
| Zamia neurophyllidia D.W.Stev. | V (Isthmus): Costa Rica, Nicaragua, Panama | Least concern |
| Zamia obliqua A.Braun | V (Isthmus): Colombia | Least concern |
| Zamia onan-reyesii C.Nelson & Sandoval | III-B (Mega Mexico): Honduras | Endangered |
| Zamia oreillyi C.Nelson | III-B (Mega Mexico): Honduras | Critically endangered |
| Zamia orinoquiensis Calonje, Betancur & A.Linstr.[24] | VII (East of Andes): Columbia | Not evaluated |
| Zamia paucifoliolata Calonje[25] | VI (West of Andes): Colombia | Not evaluated |
| Zamia paucijuga Wieland | III-A (Mega Mexico): Mexico (Guerrero, Jalisco, Colima, Michoacán, Nayarit, Oaxaca) | Near threatened |
| Zamia poeppigiana Mart. & Eichler | VII (East of Andes): Bolivia, Brazil (Acre) | Least concern |
| Zamia portoricensis Urb. | I (Caribbean): Puerto Rico | Endangered |
| Zamia prasina W.Bull | III-A (Mega Mexico): Belize, Mexico (Tabasco, Yucatán) | Least concern |
| Zamia pseudomonticola L.D.Gómez | V (Isthmus): Costa Rica, Panama | Least concern |
| Zamia pseudoparasitica Yates | V (Isthmus): Panama | Near threatned |
| Zamia pumila L. - guáyara / Dominican zamia | I (Caribbean): Hispaniola (Dominican Republic), possibly Cuba; possibly extirpated in Puerto Rico and Haiti | Vulnerable |
| Zamia purpurea Vovides, J.D.Rees & Vázq.Torres | III-A (Mega Mexico): Mexico (Veracruz, Oaxaca) | Critically endangered |
| Zamia pygmaea Sims | I (Caribbean): Cuba | Endangered |
| Zamia pyrophylla Calonje, D.W.Stev. & A.Lindstr. | VI (West of Andes): Colombia | Critically endangered |
| Zamia restrepoi (D.W.Stev.) A.Lindstr. | VII (Manicata): Colombia | Critically endangered |
| Zamia roezlii Regel ex Linden | VI (West of Andes): Colombia, Ecuador | Least concern |
| Zamia sandovalii C.Nelson | III-B (Mega Mexico): Honduras | Endangered |
| Zamia sinuensis Calonje & Castro[26] | VII (Manicata): Columbia | Not evaluated |
| Zamia skinneri Warsz. ex A.Dietr. | V (Isthmus): Panama | Endangered |
| Zamia soconuscensis Schutzman, Vovides & Dehgan | IV: Mexico (Chiapas) | Endangered |
| Zamia spartea A.DC. in A.P.de Candolle | III-A (Mega Mexico): Mexico (Oaxaca) | Critically endangered |
| Zamia splendens Schutzman[27] | III-A (Mega Mexico): Mexico (Chiapas, Tabasco, Veracruz) | Endangered |
| Zamia standleyi Schutzman | III-B (Mega Mexico): Guatemala, Honduras | Not evaluated |
| Zamia stenophyllidia Nicolalde-Morejón, Martínez-Domínguez & Stevenson[28] | (unknown): Mexico (Michoacán) | Not evaluated |
| Zamia stevensonii A.S.Taylor & Holzman | V (Isthmus): Panama | Endangered |
| Zamia stricta Miq. | I (Caribbean): Cuba | Vulnerable |
| Zamia tolimensis Calonje, H.E.Esquivel & D.W.Stev | VII (East of Andes): Colombia | Endangered |
| Zamia tuerckheimii Donn.Sm. | III-B (Mega Mexico) Guatemala | Near threatened |
| Zamia ulei Dammer | VII (East of Andes): Brazil, Colombia, Ecuador, Peru, Venezuela | Least concern |
| Zamia urep B.Walln. | VII (East of Andes): Peru | Endangered |
| Zamia variegata Warsz. | III-A (Mega Mexico): Belize, Guatemala, Mexico (Chiapas) | Endangered |
| Zamia vazquezii D.W.Stev., Sabato & De Luca | II (Fischeri): Mexico (Veracruz) | Critically endangered |
| Zamia wallisii H.J.Veitch | VI (West of Andes): Colombia | Critically endangered |
References
[edit]- ^ "Genus: Zamia L." Germplasm Resources Information Network. United States Department of Agriculture. 2007-10-05. Archived from the original on 2009-08-27. Retrieved 2011-01-12.
- ^ a b Kew World Checklist of Selected Plant Families
- ^ Hill, K.D. & Stevenson, D.W. (1999). A world list of Cycads, 1999. Excelsa 19: 67-72.
- ^ Flora of North America, vol 2, Zamia integrifolia Linnaeus f. in Aiton, Hort. Kew. 3: 478. 1789.
- ^ Standley, P. C. & J. A. Steyermark. 1958. Cycadaceae. In Standley, P.C. & Steyermark, J.A. (Eds), Flora of Guatemala - Part I. Fieldiana, Bot. 24(1): 11–20.
- ^ Monteza-Moreno, Claudio M.; Rodriguez-Castro, Lilisbeth; Castillo-Caballero, Pedro L.; Toribio, Edgar; Saltonstall, Kristin (2022). "Arboreal camera trapping sheds light on seed dispersal of the world's only epiphytic gymnosperm: Zamia pseudoparasitica". Ecology and Evolution. 12 (3): e8769. Bibcode:2022EcoEv..12E8769M. doi:10.1002/ece3.8769. PMC 8948316. PMID 35356569.
- ^ Nicolalde-Morejón, F., A. P. Vovides & D. W. Stevenson. 2009. Taxonomic revision of Zamia in Mega-Mexico. Brittonia 61(4): 301–335.
- ^ Armstrong, W.P. (2008-04-03). "Botanical Record-Breakers (Part 1 of 2)". Wayne's Word. Archived from the original on 2010-12-19. Retrieved 2011-01-12.
- ^ 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; Smith, Stephen A.; Yi, Ting-Shuang; et al. (2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. Bibcode:2021NatPl...7.1015S. bioRxiv 10.1101/2021.03.13.435279. doi:10.1038/s41477-021-00964-4. PMID 34282286. S2CID 232282918.
- ^ Stull, Gregory W.; et al. (2021). "main.dated.supermatrix.tree.T9.tre". Figshare. doi:10.6084/m9.figshare.14547354.v1.
{{cite journal}}: Cite journal requires|journal=(help) - ^ Zonneveld, B. J. M.; Lindström, A. J. (26 May 2016). "Genome sizes for 71 species of Zamia (Cycadales: Zamiaceae) correspond to three different biogeographic regions". Nordic Journal of Botany. 34 (6): 744–751. doi:10.1111/njb.01094.
- ^ Calonje, Michael; Meerow, Alan W.; Griffith, M. Patrick; Sala-Leiva, Dayana; Vovides, Andrew P.; Ciro, Mario; Francisco-Ortega, Javier (May 2019). "A Time-Calibrated Species Tree Phylogeny of the New World Cycad Genus Zamia L. (Zamiaceae, Cycadyles)". International Journal of Plant Science. 180 (4): 294. doi:10.1086/702642.
- ^ a b Lindstrom, Anders; Nabib, Sadaf; Dong, Shanshan; Dong, Yiqing; Liu, Jiang; Calonje, Michael; Stevenson, Dennis; Zhang, Shouzhou (2024). "Transcriptome sequencing data provide a solid base to understand phylogenetic relationships, biogeography and reticulated evolution of the genus Zamia L. (Cycadales, Zamiaceae)". Annals of Botany. XX: 10 (chart), 16–17. doi:10.1093/aob/mcae065. PMID 38900840.
- ^ Eckenwalder, James E. (October 1980). "Taxonomy of the West Indian cycads". Journal of the Arnold Arboretum. 61 (4): 715–720. JSTOR 43782079.
- ^ Calonje, M.; Stevenson, Dennis; Osborne, Roy (2024). "About the World List of Cycads". World List of Cycads. Retrieved October 14, 2024.
- ^ Calonje, Michael; Meerow, Alan W.; Knowles, Lindy; Knowles, David; Griffith, M. Patrick; Nakamura, Kyoko; Francisco-Ortega, Javier (April 2013). "Cycad biodiversity in the Bahamas Archipelago and conservation genetics of the threatened Zamia lucayana (Zamiaceae)". Oryx. 47 (2). Introduction. doi:10.1017/S0030605312000129.
- ^ "Zamia — The Plant List". www.theplantlist.org. Retrieved 2019-06-04.
- ^ "Zamia brasiliensis". The World List of Cycads. Retrieved 15 October 2024.
- ^ Calonje, Michael; Hernanzez, Jonaton Castro; Coca, Luis Fernando; Jaramillo, Daniel; Aristzabal, Arturo (16 April 2016). "Two new species of Zamia (Zamiaceae, Cycadales) from the Magdalena-Uraba moist forests of northern Colombia". Phytotaxa. 497 (1). doi:10.11646/phytotaxa.497.1.1.
- ^ Stevenson, Dennis Wm.; Cardena Lopez, Dairon; Castano Arboleda, Nicolas (19 June 2018). "A new Zamia (Zamiaceae) from Colombia". Brittonia. 70: 364–368. doi:10.1007/s12228-018-9535-3 – via Springer Link.
- ^ "Zamia magnifica". The World List of Cycads. Retrieved 15 October 2024.
- ^ "Zamia multidentata". The World List of Cycads. Retrieved 15 October 2024.
- ^ "Zamia nana". The World List of Cycads. Retrieved 15 October 2024.
- ^ "Zamia orinoquiensis". The World List of Cycads. Retrieved 15 October 2024.
- ^ Calonje, Michael; Lopez-Gallego, Cristina; Castro, Jonaton (28 December 2018). "Zamia paucifoliolata, a new species of Zamia (Zamiaceae, Cycadales) from Valle de Cauca, Colombia". Phytotaxa. 385 (2): 364–368 – via Biotaxa.
- ^ Calonje, Michael; Hernanzez, Jonaton Castro; Coca, Luis Fernando; Jaramillo, Daniel; Aristzabal, Arturo (16 April 2016). "Two new species of Zamia (Zamiaceae, Cycadales) from the Magdalena-Uraba moist forests of northern Colombia". Phytotaxa. 497 (1). doi:10.11646/phytotaxa.497.1.1.
- ^ "Zamia splendens". The World List of Cycads. Retrieved 15 October 2024.
- ^ Nicolalde-Morejón, Fernando; Martínez-Domínguez, Lilí; Stevensob, Dennis Wm.; Vergara-Silva, Francisco (September 2019). "Disentangling the identity of Zamia from a Mexican Pacific seaboard, with a description of a new species". Nordic Journal of Botany. 37 (9). doi:10.1111/njb.02430.
External links
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