Jump to content

Zamia

From Wikipedia, the free encyclopedia
For the three species commonly called Zamia that are endemic to Western Australia, see Macrozamia riedlei, Macrozamia fraseri, and Macrozamia dyeri.

Zamia
Zamia furfuracea
Scientific classification Edit this 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]
  • Aulacophyllum Regel
  • Chigua D.W.Stev.
  • Palmifolium Kuntze
  • Palma-filix Adans.

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]
Zamia furfuracea leaves

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). Z. picta is even more distinctive, being the only truly variegated cycad (having whitish/yellow speckles on the leaves).[7]

Like all Zamiaceae, Zamia plants have "coralloid" (coral-shaped) roots which host nitrogen-fixing cyanobacteria. Stems are 3 to 25 centimetres (1.2 to 9.8 in) in diameter, and when arborial, up to 5 metres (16 ft) tall. Leaves vary from 2 to up to 15 in number, and have an even number (5 to 60 pairs) of leaflets. petioles (leaf stalks) and rachis (leaf axis, to which leaflets are attached) may be smooth, or have prickles on the petiole and lower part of the rachis. Leaflets are linear to ovate and may be directly attached to the rachis, or may be on short petiolules. The edges of leaflets may be smooth, or may be toothed partly or around the whole leaflet.[8] The morphology of leaflets is highly varied between even closely related species of Zamia, and within species and even within populations of a species. Studies have found that the amount of sun a plant is exposed to is responsible for differences in leaflet length, width, surface area, width ratio, shape, and density and thickness of leaflets. There is also significant morphological variation in leaflets between male and female plants in some species.[9][10]

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.[11]

It was long believed that Zamia plants, like all cycads, relied completely on wind pollination. In the 1980s it was discovered that at least some Zamia species were pollinated by beetles of the Pharaxonotha and Rhopalotria genera. Further studies have found that pollination by beetles is widespread in Zamia and other cycads.[12]

Seed-dispersal in Zamia is poorly documented for most species, but there are reports of birds and/or small to medium-sized mammals dispersing the seeds of a few Zamia species.[12]

Habitats

[edit]

With one exception, Zamia species are found only in the Neotropical realm in the Americas. The exception is Z. integrifolia, which has a range that extends into the Nearctic realm in northern Florida, and formerly into the southeastern corner of Georgia.[13]

Toxicity

[edit]

The primary toxin in Zamia plants is cycasin, a carcinogenic and neurotoxic glucoside. Other glucoside toxins present include macrozamin (de) and several neocycasins.[14] BMAA, a neurotoxin that is produced by cyanobacteria living in roots of the plants, is also present in most Zamia plants.[15]

Many of the Zamia species are, or have been, gathered to process the stem and/or seeds into starch for use as food or laundry starch, or for use as traditional medicine. Since almost the entire plant is very toxic, the starch must be grated and repeatedly washed to remove the toxins and make the starch edible. Only the sarcotesta, the pulpy covering of the seeds, is relatively free of toxins.[16]

Consumption of cycads by livestock has resulted in two forms of cycad toxicosis. Hepato-gastrointestinal toxicosis results from damage to the liver and gastrointestinal tracts of affected animals causing depression, anorexia, and weight loss. Neurologic toxicosis, known as Zamia staggers, is the result of damage to brain, spinal cord, and dorsal root ganglia tissue causing weight loss, swaying of hind quarters, and weakness and other defects in rear limbs.[17]

Herbivory

[edit]

The presence of toxins protects Zamia plants from most herbivores, but caterpillars of the genus Eumaeus are obligate herbivores of cycads, including Zamia species. Eumaeus caterpillars have aposematic coloration, being bright red with yellow or white bands, and adults have red elements on their body and wings. The caterpillars sequester and retain the toxins consumed from Zamia plant tissue, rendering all stages of Eumaeus distasteful to predators. With few exceptions, Zamia species host only one species of Eumaeus caterpillars each.[18]

The known host dependencies of Eumaeus species on Zamia species include: Species in the Florida/Caribbean clade, or Z. pumila species complex, are hosts for caterpillars of Eumaeus atala. Species in the Fisheri clade, and Z. cremnophila of the Mega-Mexico A sub-clade, are hosts for caterpillars of E. childrenae. Some species in the Z. loddigesii species complex of the Mega-Mexico A sub-clade are hosts for caterpillars of E. toxea. Many species in the South American clade west of the Andes are hosts for caterpillars of E. godartii. Several species in the South American clade east of the Andes are hosts for caterpillars of E. minyas. One Zamia species found east of the Andes, Z. poeppigiana, is host for caterpillars of both E. minyas and E. toxana. Another Zamia species east of the Andes, Z. amazonum, also hosts caterpillars of E. toxana.[19] The use of other Zamia species as hosts for Eumaeus species had not been established as of 2023.[18]

Caterpillars of the moths Seirarctia echo and Anatrachyntis badia have also been observed on Zamia integrifolia, and caterpillars of an unknown species of a Blastobasidae moth have been observed on Zamia pumila.[20]

Conservation

[edit]

Most species of Zamia are prized as ornamental plants, but are difficult to propagate and slow growing. As a result, Zamia plants are often over-collected, threatening to wipe out populations or whole species. In order to protect wild populations, scientists working with Zamia species no longer publish the exact locations of populations.[21]

Karyology

[edit]

Zamia is the only genus in the cycads that has variation in karyotypes between species, specifically in the chromosome number. Diploid chromosome numbers in Zamia include 2n=16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, and 28. Some Zamia species have varying numbers of diploid chromosomes within the species. The odd diploid chromosome numbers occur in species with more than one chromosome number.[22][23]

Phylogeny

[edit]

Despite the ancient history of cycads, species diversity in Zamia is geologically recent. Calonje et al. found a stem age for Zamia of 68.28 million years ago (mya), and a crown age of 9.54 mya.[24]

Geographic groups

[edit]

As early as the 1930s, authors recognized three biogeographic groups in Zamia, Caribbean, Mesoamerican, and Central and South America. In the 21st century, phylogenies of Zamia based on molecular phylogenetic analyses have found stronger correlation with geographic regions than with morphological features.[25]

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 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.[26]

Zamia tree based on DNA[27]
Zamia

Caribbean and Florida 8 sp.

Mesoamerica 21 sp.

Zamia soconuscensis

Isthmus 16 sp.

South America 28 sp.

Calonje, et al. (2019) analyzed the DNA from 70 species of Zamia, finding support for four geographically distinct clades (plus a single isolated species). A clade including the species found on the Caribbean islands and in Florida is sister to the rest of the genus. The species of the Caribbean clade have diverged within the last 1.9 million years. The Mesoamerica clade includes all species found in Mesoamerica (north of Nicaragua), except for the single species Z. soconuscensis. It has a divergence age of 5.79 mya. The Isthmus clade includes species found in southernmost Nicaragua, Costa Rica and Panama, and has a divergence age of 2.35 mya. The species in South America form another clade, which is sister to the Isthmus clade. It has a divergence age of 2.62 mya.[27]

Zamia tree based on transcriptomes[28]
Zamia
I

Caribbean islands 7 sp.

Florida Zamia integrifolia

II

Fischeri clade 3 sp.

III
III‑A

Mega Mexico A 14 sp.

III‑B

Mega Mexico B 7 sp.

IV

Zamia soconuscensis

V

Isthmus 15 sp.

VI

West of the Andes 12 sp.

VII

Northern Colombia

(Manicata clade) 4 sp.

East of the Andes 13 sp.

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 eight 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 Clade I, with the group of Clade I and Clade II being sister to the rest of the genus. Clade III (Mega Mexico) is divided into the sub-clades III-A and III-B. Clade III-A includes 14 species found in Mexico and northern Central America. Clade III-B consists of seven species found in Honduras, Guatemala, and Belize. Clade IV consists of the single species Z. soconuscensis found in cloud forests in Chiapas state in Mexico. Clade V (the Isthmus clade) includes 15 species found in southern Nicaragua, 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.[28]

Clades and species complexes

[edit]

Fischeri clade

[edit]

The Fischeri clade consists of three closely related species of Zamia found near the Gulf coast of central Mexico. The three species, Z. fischeri, Z. inermis, and Z. vazquezii, share various morphological features, including the near or total absence of prickles on leaf stalks. The analysis of DNA by Calonje, et al. found strong support for the Fischeri clade as sister to all of the mainland Zamia species (i.e., everything except the Caribbean and Florida clade.[29] The analysis of transcriptomes by Lindstrom et al. found strong support for the Fischeri clade as sister to the Caribbean-Florida clade, with the combined Caribbean-Florida and Fischeri clade sister to the rest of Zamia. The Fischeri and Caribbean-Florida clades share several morphological traits, including the lack or near-lack of prickles on leaf stalks and simlarities in reproductive characters. The genome sizes of the species in the Fischeri clade are among the smallest in Zamia.[30]

Manicata clade

[edit]

In 1996, Caputo et al. described a "Manicata clade" consisting of Zamia manicata, Z. cunaria, Z. obliqua, and Z. iepetiensis.[31] Later molecular phylogenetic studies have produced a different definition of the clade, with Z. manicata being the only species included in the new definition. The Manicata clade is a group of closely related species found in northern Colombia. It consists of Z. manicata, Z. disodon, Z. melanorrhacis, Z. restrepoi, Z. imbricata, and Z. sinuensis. Z. manicata was described in 1876. The other species in the clade have been described more recently, with Z. restrepoi described in 1990 (as Chigua restrepoi, reclassified as Z. restrepoi in 2009), Z. disodon and Z. melanorrhacis in 2001, and Z. imbricata and Z. sinuensis in 2021. The monophyly of the clade is strongly supported by molecular phylogenetic studies. Calonje, et al. (2019) found Z. manicata, Z. disodon, Z. melanorrhacis, and Z. restrepoi to form a clade.[32] Lindstrom et al. (2024) found Z. manicata, Z. disodon, Z. restrepoi, and Z. sinuensis to form a clade.[33] While Z. imbricata was not included in either analysis, it shares several morphological features with the rest of the clade, including subterranean or semi-subterranean stems, strongly toothed margins on leaflets, strobili (cones) on very long stalks, small seeds with very thin sarcotesta (coats), and very small pollen (male) cones. Leaf morphology varies strongly among the members of the clade.[34]

Zamia loddigesii species complex

[edit]
Zamia loddigesii species complex tree based on DNA[29]
Zamia

A genetic study by Cafasso et al. published in 2009 found a basis for a Zamia loddigesii species complex in the presence of the same four discrete clusters of DNA repeats in Z. loddigesii, Z. furfuracea, Z. picta, Z. paucijuga, Z. polymorpha, Z. purpurea, Z. spartea, Z. splendens, and Z. sylvatica.[35] (After the study was published, Z. sylvatica was reclassified as a synonym of Z. loddigesii,[36] Z. picta was reclassified as a synonym of Z. variegata,[37] and Z. polymorpha was reclassified as a synonym of Z. prasina.[38]) A 2019 study of DNA in Zamia found a clade of 12 species that includes all of the members of the species complex defined by the Cafasso et al. study. (Z. lacandona, Z. grijalvensis, and Z. herrerae were not included in the study by Cafasso et al., but Z. cremnophila was.)[29] Gutiérrez-Ortega et al. call this the "Mexican" clade.[23]

Another 2019 study found a large variation in chromosome number within a few Zamia species, including four in the species complex, Z. loddigesii (2n=17–18, 24–27), Z. paucijuga (2n=19, 23–28), Z. prasina (2n=22–27), and Z. lacandona (2n=16–18). Z. variegata has a smaller intraspecific variation in chromosome number (2n=21–22). Two species in the Z. loddigesii complex, including Z. furfuracea and Z. spartea, have a chromosome number of 2n=18. Z. cremnophila and Z. purpurea have a chromosome number of 2n=16, while Z. herrerae has one of 2n=24.[39]

Zamia pumila species complex

[edit]
Zamia pumila species complex tree based on transcriptomes[28]
Zamia

All of the species in the Caribbean plus Florida grouping/clade are also in the Zamia pumila species complex. The classification of the populations in the Greater Antilles, Bahamas, and Florida has been controversial. In 1980, Eckenwalder included all the Zamia populations in the Caribbean and Florida in the single species Z. pumila, incorporating 27 previously described species (not all of which were valid or accepted at the time) into the subspecies Z. pumila subsp. pumila, and five such species from Cuba into the subspecies Z. pumila subsp. pygmaea.[40] Eckenwalder's classification is no longer generally accepted, and a monophyletic species complex consisting of nine species is now accepted, including Z. pumila, seven other 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.[41][42] The 2009 study by Cafasso et al. found support for the species complex in the presence of the same four discrete clusters of DNA repeats in Z. integrifolia, Z. portoricensis, and Z. pumila. The other species in this complex were not examined in that study.[43]

Zamia skinneri species complex

[edit]
Z. skinneri species complex tree based on transcriptomes[28]
Zamia

Zamia skinneri has been regarded as a highly variable species. Z. neurophyllidia was described as a new species in 1993, based on a population of what had been regarded as a dwarf form of Z. skinneri. A study published in 2004 proposed that Z. neurophyllidia and Z. skinneri were a "hybrid species complex", and noted that Z. skinneri included several morphologically distinct populations. In 2008, Taylor B. et al. described three sub-populations of Z. skinneri as Z. hamannii, Z. imperialis, and Z. nesophila.[44] All five of the species in this complex are found in Bocas del Toro Province in Panama, at least three of the species are endemic to that province, and all of them have plicate leaves, a trait that occurs elsewhere in Zamia only in Z. dressleri in Colon and Guna Yala (formerly San Blas) provinces in Panama, and in Z. roezlii and Z. wallisii in Colombia. Taylor B. et al. (2012) suggest that Z. skinneri is the central species of the complex, and that the other species have evolved rapidly from Z. skinneri on the periphery of its range due to geographic or other isolation.[45]

Chigua

[edit]

Chigua was described as a new genus with two species in Zamiaceae in 1990, but was reclassified as Zamia restrepoi in 2009.[46]

Species

[edit]

The delimitation of species in Zamia has been problematic. Early taxonomy of the genus was based primarily on leaflet morphology, including leaflet shape, vein number, leaflets per leaf, and apex shape. Based on differences in leaflet morphology, named species proliferated, with 128 species of Zamia having been named by the 1980s, often based on a single specimen of unknown type site. Many of those proposed species have been moved to other genera or synonymized with accepted Zamia species. By 1985, moreover, it had been demonstrated that leaflet morphology was influenced by environment and thus not necessarily useful for distinguishing species.[47] It is now recognized that reproductive structures, especially female strobili (cones), are essential to distinguishing species of Zamia.[48]

The number of recognized species in Zamia has more than doubled since the 1980s. There were approximately 40 species recognized in 1988,[49] 58 species in 2008,[50] 76 in 2016,[51] and 80 in 2019.[52] As of 2025, the World List of Cycads lists 89 accepted species of Zamia,[53] the Royal Botanic Gardens, Kew, recognizes 84,[54] and World Flora Online recognizes 89.[55]

Accepted species[α] Clade:[28] 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 (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. 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[β] Calonje & J.Castro VII (Manicata): Colombia[10] 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. 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 lawsoniana[γ]' Dye (unknown): Mexico (Chiapas, Oaxaca, Tabasco, Veracruz) Not evaluated
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 (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[δ] Pérez-Farr., Gutt-Ortega & Calonje (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. VII (Manicata): Colombia[10] Endangered
Zamia montana A.Braun VI (West of Andes): Colombia Critically endangered
Zamia monticola Chamb. III-B (Mega Mexico): Guatemala Critically endangered
Zamia multidentata[ε] Calonje, Segalla & R.S.Pimenta (unknown): Brazil (Acre) Not evaluated
Zamia muricata Willd. VII (East of Andes): Colombia, Venezuela Least concern
Zamia nana[ζ] A.Lindstr., Calonje, D.W.Stev. & A.S.Taylor 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 oligodonta E.Calderón & D.E.Stev. (unknown): Colombia Critically endangered
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[η] Calonje, Betancur & A.Linstr. VII (East of Andes): Colombia Not evaluated
Zamia paucifoliolata Calonje 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 threatened
Zamia pumila L. 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[θ] Calonje & J.Castro VII (Manicata): Colombia 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[ι] Schutzman 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 (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 verschaffeltii Miq. (unknown): Mexico Not evaluated[κ]
Zamia wallisii H.J.Veitch VI (West of Andes): Colombia Critically endangered

Species table notes

[edit]
  1. ^ Almost all of the species in this table are accepted as valid in at least four of the following databases, including The World List of Cycads (WLC),[56] the USDA-ARS Germplasm Resources Information Network (GRIN),[57] the Kew Gardens Plants of the World Online (Kew),[58] the World Flora Online (WFO).[59] and Tropicos.[60] A few species accepted in only two or three of the five databases are noted individually.
  2. ^ Z. imbricata is accepted in WLC, Kew, and Tropicos.
  3. ^ Z. lawsoniana is accepted in WLC, WFO, and Tropicos.
  4. ^ Z. magnifica is accepted in WLC, Kew, and Tropicos.
  5. ^ Z. multidentata is accepted in WLC, Kew, and Tropicos.
  6. ^ Z. nana is accepted in WLC, Kew, and Tropicos.
  7. ^ Z. orinoquiensis is accepted in WLC and Kew, and listed as "unchecked" in WFO.
  8. ^ Z. sinuensis is accepted in WLC, Kew, and Tropicos.
  9. ^ Z. splendens is accepted in WLC, WFO, and Tropicos.
  10. ^ Z. verschaffeltii may be extinct.[61]

References

[edit]
  1. ^ "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.
  2. ^ a b Kew World Checklist of Selected Plant Families
  3. ^ Hill, K.D. & Stevenson, D.W. (1999). A world list of Cycads, 1999. Excelsa 19: 67-72.
  4. ^ Flora of North America, vol 2, Zamia integrifolia Linnaeus f. in Aiton, Hort. Kew. 3: 478. 1789.
  5. ^ 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.
  6. ^ 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.
  7. ^ Nicolalde-Morejón & VovidesStevenson 2009, pp. 301–335.
  8. ^ Stevenson 2004, p. 197, 202.
  9. ^ Lindstrom 2009, pp. 269–270.
  10. ^ a b c Calonje et al. 2021, p. 1.
  11. ^ 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.
  12. ^ a b Segalla et al. 2019, Mutualistic Interaction: Plant-Pollinators.
  13. ^ Calonje et al. 2019, p. 286.
  14. ^ Schneider, Dietrich; Wink, Michael; Sporer, Frank; Lounibos, Philip (July 2002). "Cycads: their evolution, toxins, herbivores and insect pollinators". Naturwissenschaften. 89 (7): 283. Bibcode:2002NW.....89..281S. doi:10.1007/s00114-002-0330-2. PMID 12216856 – via Springer Link.
  15. ^ Holtcamp, Wendee (1 March 2012). "The Emerging Science of BMAA: Do Cyanobacteria Contribute to Neurodegenerative Disease?". Environmental Health Perspectives. 120 (3): a110 – a116. doi:10.1289/ehp.120-a110. PMC 3295368. PMID 22382274.
  16. ^ Stevenson, Dennis Wm. (1991). "The Zamiaceae of the Southeastern United States". Journal of the Arnold Arboretum, Supplementrary Series. 1: 376, 381. doi:10.5962/p.315947. JSTOR 43782788.
  17. ^ Reams, Rachel Y.; Janovitz, Evan B.; Robinson, Farrel R.; Sullivan, John M.; Casanova, Carlos Rivera; Más, Edwin (July 1993). "Cycad (Zamia puertoriquensis) Toxicosis in a Group of Dairy Heifers in Puerto Rico". Journal of Veterinary Diagnostic Investigation. 5 (3): 488–494. doi:10.1177/104063879300500337. ISSN 1040-6387. PMID 8373871.
  18. ^ a b Sierra‐Botero et al. 2023, p. 2.
  19. ^ Sierra‐Botero et al. 2023, p. 7.
  20. ^ Whitaker & Salzman 2020, p. 1864.
  21. ^ Stevenson, Dennis Wm. (January 1993). "The Zamiaceae in Panama with Comments on Phytogeography and Species Relationships". Brittonia. 45 (1): 2. doi:10.2307/2806850.
  22. ^ {{Cite journal |last=Caputo |first=Paolo |last2=Cozzolino |first2=Salvatore |last3=Gaudio |first3=Lucinao |last4=Moretti |first4=Aldo |last5=Stevenson |first5=Dennis W. |date=1996 |title=Karyology and Phylogeny of Some Mesoamerican Species of Zamia (Zamiaceae) |url=https://www.jstor.org/stable/2446105?seq=1 |journal=American Journal of Botany |volume=83 |issue=11 |pages=1513–1520 |doi=10.2307/2446105 |issn=0002-9122 |jstor=}2446105}
  23. ^ a b Gutiérrez-Ortega et al. 2023, pp. 111–112.
  24. ^ Calonje et al. 2019, p. 294.
  25. ^ Calonje et al. 2019, pp. 287–288.
  26. ^ Zonneveld & Lindström 2016, p. 750.
  27. ^ a b Calonje et al. 2019, pp. 287–294.
  28. ^ a b c d e Lindstrom et al. 2024, pp. 10 (chart), 16–17.
  29. ^ a b c Calonje et al. 2019, p. 301.
  30. ^ Lindstrom et al. 2024, p. 762.
  31. ^ Caputo, Paolo; Cozzolino, Salvatore; Gaudio, Lucinao; Moretti, Aldo; Stevenson, Dennis W. (1996). "Karyology and Phylogeny of Some Mesoamerican Species of Zamia (Zamiaceae)". American Journal of Botany. 83 (11): 1519. doi:10.2307/2446105. ISSN 0002-9122. JSTOR 2446105.
  32. ^ Calonje et al. 2019, pp. 296 (tree).
  33. ^ Lindstrom et al. 2024, pp. 10 (tree), 19.
  34. ^ Calonje et al. 2021, pp. 1, 5, 9.
  35. ^ Cafasso et al. 2009, p. 32.
  36. ^ "Zamia sylvatica". The World List of Cycads. Retrieved 2025-03-06.
  37. ^ "Zamia picta". The World List of Cycads. Retrieved 2025-03-07.
  38. ^ "Zamia polymorpha". The World List of Cycads. Retrieved 2025-03-07.
  39. ^ Rastogi, Shubhi; Ohri, Deepak (2020). "Karyotype evolution in cycads". The Nucleus. 63 (2): 132, 135. doi:10.1007/s13237-019-00302-2. ISSN 0029-568X.
  40. ^ Eckenwalder 1980, pp. 715–720.
  41. ^ Calonje, M.; Stevenson, Dennis; Osborne, Roy (2024). "About the World List of Cycads". World List of Cycads. Retrieved October 14, 2024.
  42. ^ 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.
  43. ^ Cafasso et al. 2009, p. 33.
  44. ^ Taylor B., Alberto S.; Haynes, J. L.; Holzman, G. (2008). "Taxonomical, nomenclatural and biogeographical revelations in the Zamia skinneri complex of Central America (Cycadales: Zamiaceae)" (PDF). Botanical Journal of the Linnean Society. 158 (3): 403–404, 408–424. doi:10.1111/j.1095-8339.2008.00886.x.
  45. ^ Taylor B., Alberto S.; Haynes, Jody L.; Stevenson, Dennis W.; Holzman, Gregory; Mendieta, Jorge (2012). "Biogeographic Insights in Central American Cycad Biology" (PDF). In Stevens, Lawrence (ed.). Global Advances in Biogeography. Rejika, Croatia: InTechOpen. pp. 73–98. ISBN 978-9535104544.
  46. ^ Lindstrom et al. 2024, p. 17.
  47. ^ Schutzman, Vovides & Dehgan 1988, pp. 358–359.
  48. ^ Nicolalde-Morejón, Vovides & Stevenson 2009, p. 303.
  49. ^ Schutzman, Vovides & Dehgan 1988, p. 347.
  50. ^ Nicolalde-Morejón et al. 2008, p. 38.
  51. ^ Zonneveld & Lindström 2016, p. 744.
  52. ^ Segalla et al. 2019, Introduction.
  53. ^ "Zamia". World List of Cycads. Retrieved 2025-02-28.
  54. ^ "Zamia". Plants of the World Online. Retrieved 2025-03-12.
  55. ^ "Zamia L." World Flora Online. Archived from the original on 2023-01-05. Retrieved 2025-03-12.
  56. ^ "The World List of Cycads, online edition". www.cycadlist.org. Retrieved 21 November 2024.
  57. ^ "USDA-ARS Germplasm Resources Information Network (GRIN)". www.ars-grin.gov. Retrieved 2024-12-03.
  58. ^ "Plants of the World Online". Royal Botanic Gardens, Kew. Retrieved 21 November 2024.
  59. ^ "An Online Flora of All Known Plants". The World Flora Online. 2024. Retrieved 21 November 2024.
  60. ^ "Tropicos". tropicos.org. Retrieved 2024-12-03.
  61. ^ Nicolalde-Morejón et al. 2008, pp. 45–46.

Sources

[edit]
[edit]
Wikimedia Commons has media related to Zamia.
Wikispecies has information related to Zamia.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Zamia&oldid=1280225031"