Zamia nesophila

Zamia nesophila is a shrubby to tree-like cycad of the islands of the Bocas del Toro archipelago, in north-western Panama, where most plants grow in beach sand just above the high-tide line. Long included in Zamia skinneri, this species of the genus Zamia was described in 2008. The IUCN rates it Critically Endangered (2022 assessment): tourist and residential development of the coast has destroyed a large part of its populations.

Key facts about Zamia nesophila

  • Accepted name: Zamia nesophila A.S.Taylor, J.L.Haynes & Holzman (2008), accepted by The World List of Cycads (version 2026.08.15), by GBIF and by the World Checklist of Vascular Plants, on which POWO is based; POWO could not be consulted directly
  • Former names and synonyms: no synonym; the island populations were previously placed in Zamia skinneri
  • Family: Zamiaceae, order Cycadales
  • Common names: no English name; “guade teet”, a local name in the Bocas del Toro archipelago (original description)
  • Natural range: Panama, province of Bocas del Toro (islands of the archipelago), from sea level to low hills
  • IUCN status: Critically Endangered (CR), criteria A2acde+4acde; B1ab(i,ii,iii,iv,v), 2022 assessment
  • CITES and other protection: CITES Appendix II (whole genus, since 1977; seeds exempt under annotation #4); Annex B in the European Union
  • Documented hardiness: young plants came through the winter of 2014–2015 in the ground, in shade, on the Mediterranean coast west of Marseille, where the nearest weather station recorded −3.0 °C (26.6 °F); defoliated by about −2 to −2.5 °C (28.4 to 27.5 °F) in Greece, according to a second-hand report (see Cold hardiness)
  • Light and soil: calcareous beach sand, salt spray; partial shade in cultivation, though it takes more sun than its close relatives
  • Habit and adult size: trunk up to 2.8 m tall and 6 to 24 cm across; up to 20 leaves 116 to 239 cm long, with 5 to 12 pairs of leaflets (original description)
  • In the UK, US and Australia: Montgomery Botanical Center, Fairchild and nine other American gardens (US); Royal Botanic Garden Sydney (Australia); no British record
  • Toxicity: no data specific to the species; treat it as poisonous, like all cycads (see Toxicity and precautions)

How to recognise Zamia nesophila

It is a large Zamia with leaflets pleated between the veins, leaves that emerge bright green, and a seashore habitat. The original description by Alberto Taylor, Jody Haynes and Greg Holzman (2008) gives:

  • Stem: a shrub or small tree with a trunk up to 2.8 m tall and 6 to 24 cm in diameter, usually solitary, sometimes branched near the base or the apex, especially after damage.
  • Leaves: up to 20 per crown (11 on average), 116 to 239 cm long, with 5 to 12 pairs of leaflets; petiole 27 to 113 cm, medium green.
  • Leaflets: elliptic, acuminate, pleated between the veins on the upper surface, with toothed margins mainly in the upper third; apical leaflets 19 to 39 cm by 7 to 15 cm, median leaflets 25 to 38 cm by 6.5 to 10.5 cm, basal leaflets 23.4 to 35.6 cm by 4 to 11.7 cm.
  • New leaves: bright green and hairless, maturing glossy medium green.
  • Male cones: conical-cylindrical, 8 to 16 cm long and 2 to 2.5 cm across, single or in groups of 2 to 5, covered in reddish-golden to brownish-yellow felt, on a peduncle 4.5 to 12 cm long; their protruding, down-turned scales set them apart from the cones of the other plicate-leaved Zamia of Panama.
  • Female cones: usually one per plant, 15 to 39 cm long and 4.5 to 10.4 cm across, yellow-brown to tan when emerging, maturing green or greyish green; cylindrical to cylindrical-globose, on a peduncle up to 15 cm long that often makes them hang down at maturity.
  • Seeds: ovoid to globose, 1.9 to 2.8 cm long and 1.1 to 1.8 cm across, with a bright red fleshy coat; 400 or more per cone.

According to its authors, Zamia nesophila differs from Zamia skinneri in its shorter, narrower and more numerous leaflets, its male cones with protruding, down-turned scales, and its cylindrical female cones on longer stalks, often pendent at maturity. The first leaf of seedlings usually carries two fairly broad, ovate leaflets with pointed tips (Taylor and colleagues, 2008). A French grower who raised many seedlings adds that young plants then keep no more than four leaflets per leaf until the stem is 3 to 4 cm across.

Jay Vannini (2019), a naturalist who has seen it in habitat, describes it as big to giant, green-flushing and very salt tolerant, and as the tallest of the Panamanian “groove-leaf” Zamia, with trunks up to about 3.1 m (10 ft).

Hybrids

No hybrid has been described. A French amateur who had observed the species in Panama considered it “very likely”, in 2015, that Zamia nesophila hybridises with Zamia neurophyllidia on the mainland; no publication confirms this.

Possible confusion

Zamia nesophila belongs to the Zamia skinneri complex, a group of Central American species with pleated leaflets, almost all of them present in the Bocas del Toro region. The colour of the new leaves is the simplest guide:

  • Zamia skinneri and Zamia neurophyllidia: new leaves green as well, but these grow on the mainland, in forest, and not on island beaches;
  • Zamia hamannii, the other island species of Bocas del Toro: new leaves pinkish red to orange, covered in silvery down;
  • Zamia imperialis, of central Panama: new leaves brown to reddish, large and widely spaced leaflets.

Taxonomy

Zamia nesophila is an accepted name according to The World List of Cycads, GBIF and the World Checklist of Vascular Plants, on which POWO is based (read on GBIF, version of 4 June 2026); POWO itself could not be consulted. The species was described in 2008 by Alberto S. Taylor Blake, of the University of Panama, Jody L. Haynes and Greg Holzman, in a revision of the Zamia skinneri complex published in the Botanical Journal of the Linnean Society (volume 158, pages 399 to 429). That work re-evaluated Zamia skinneri and Zamia neurophyllidia and described three new species: Zamia hamannii, Zamia imperialis and Zamia nesophila.

The island populations had until then been included in Zamia skinneri, a very variable species as then understood. They were separated on their habitat, range, new-leaf colour and chemical differences, according to the summary given in a 2010 revision of the Zamia of Costa Rica. For the reader, this means that older literature and plants sold before 2008 as Zamia skinneri from the islands of Bocas del Toro may be Zamia nesophila.

In 2012 Taylor and colleagues suggested that Zamia nesophila and Zamia hamannii diverged by isolation on the islands of the archipelago, as sea levels rose and fell, perhaps within the last 5,000 to 10,000 years; they call them “incipient species”. This is a hypothesis, not a result of phylogenetic analysis. A plant from Bocas del Toro was included in the transcriptome phylogeny of the genus published by Lindstrom and colleagues (2024), whose text does not discuss its position.

Etymology and history

The epithet nesophila comes from the Greek nêsos, “island”, and philos, “loving”: the authors chose it for the species’ “propensity for an insular existence” (Taylor and colleagues, 2008; Haynes, 2022, as quoted by The World List of Cycads).

The type was collected on 15 September 2005 in the province of Bocas del Toro, on loose sand near the shore, practically at sea level, in herbaceous and mostly shrubby vegetation with a few trees, by Alberto Taylor, Jody Haynes and Greg Holzman (no. ASTB05-BT1). The holotype is kept in the herbarium of the University of Panama (PMA), with isotypes at the Missouri Botanical Garden (MO), the New York Botanical Garden (NY), the Smithsonian Institution (US) and Xalapa (XAL).

The species was in collections before it had a name: the online catalogue of Montgomery Botanical Center, in Florida, records accessions from 2001 onwards, plus seven under “Zamia aff. nesophila”, five of them registered in 2000. In Europe it spread quickly among amateurs. A French grower bought seeds in 2010; at the end of 2012, back from a trip to Panama, he gave about a hundred to members of the Fous de palmiers forum, then 200 more in February 2015. The sowing reports published at the time, from Brittany to Provence and as far as French Guiana, make up most of what is known about its cultivation.

Common names

No English name is attested. In the Bocas del Toro archipelago the plant is known locally as “guade teet” (Taylor and colleagues, 2008); the original description does not say which language the name belongs to.

In the wild

Zamia nesophila is endemic to a few islands of the Bocas del Toro archipelago, on the Caribbean coast of north-western Panama. Its main habitat is basic, humus-covered calcareous sand among beach vegetation: the largest populations live just above the high-water mark, exposed to salt spray and occasional flooding by seawater. A secondary habitat exists on acidic, humus-rich soils in the forest of the low inland hills of some islands. The climate is tropical, with rain spread fairly evenly through the year; at certain seasons strong winds raise a heavy sea that blows spray and pushes seawater into the beach habitats. The authors count it among the very few tree-forming Zamia that are truly tolerant of saltwater. Away from houses, solitary-trunked plants nearly 3 m tall still grow under sea grape (Coccoloba uvifera) and coconut palms (Taylor and colleagues, 2008).

A French amateur who visited a population described it on the Fous de palmiers forum in 2015: plants only on the beach, on sand or rock, and absent as soon as one moves inland onto clay soil; a thin organic layer on the surface; only temporary immersion, since prolonged submersion suffocates the roots; very low seedling survival in the wild. He adds that the temperature there very rarely falls below 18 °C (64.4 °F). These are an amateur’s field observations, consistent with the published sources.

Its reproduction is unusual. According to Taylor and colleagues (2012), most Panamanian Zamia cone only once a year in the wild, but Zamia nesophila has been seen with pollen-shedding male cones and receptive female cones three times in the same year. The original description had already noted this, suggesting that more light on the beaches or the nearness of the sea might explain it; seeds take about twelve months to ripen, and seeds stripped of their fleshy coat have been found in the burrows of land crabs, possible dispersers (Taylor and colleagues, 2008).

In 2008 the pollen cones were known to shelter an unidentified Pharaxonotha beetle, assumed to be the pollinator. William Tang and colleagues, in their 2024 review of the Pharaxonotha living in Zamia cones in Panama, assign the beetles collected from male cones of Zamia nesophila in October 2012 to Pharaxonotha clarkorum, the same species as on Zamia neurophyllidia and Zamia hamannii; it appears to be the only beetle pollinator of these three cycads (Tang and colleagues, 2024; Skelley and colleagues, 2026).

Conservation status and regulation

Zamia nesophila is rated Critically Endangered (CR) by the IUCN under criteria A2acde+4acde; B1ab(i,ii,iii,iv,v), in an assessment by Alberto Taylor Blake, Jody Haynes and Greg Holzman published in 2022 (The World List of Cycads; category confirmed by GBIF); the Red List page itself could not be consulted. The A criteria reflect a rapid decline of the population, observed and expected; the B criteria, a very small range that is shrinking.

The original description details the destruction. The largest subpopulation, just above the high-water line along two white-sand beaches of about 0.8 km each, on the windward coast of one island, numbered perhaps 15,000 plants before October 2007 and dominated the undergrowth; a developer has since cleared plots along the beach and destroyed 80 to 90% of the plants. Another subpopulation on the same island, which may have held several tens of thousands of plants, had been almost entirely eliminated a few years earlier, partly with herbicides, during the building of a hotel complex; a few plants were regrowing in 2008. Other populations, reduced to a few individuals, were almost or entirely destroyed to build beach houses. The authors were already proposing a Critically Endangered listing in 2008 (Taylor and colleagues, 2008). Vannini (2019) cites it among the Panamanian cycads whose once large populations have been decimated by development in recent decades.

Like all Zamia except Zamia restrepoi, which is in Appendix I, Zamia nesophila is covered by the listing of the genus in CITES Appendix II, in force since 4 February 1977. Annotation #4 (the version adopted at CoP20, in force on 5 March 2026) exempts seeds, pollen, seedlings and tissue cultures obtained in vitro and transported in sterile containers, and cut flowers of artificially propagated plants; international trade in the plants themselves requires an export permit. In the European Union the species falls under Annex B of Regulation (EC) No 338/97, whose current list is set by Regulation (EU) 2026/1383, applicable since 29 June 2026. Species+, the official CITES database, names the species and confirms this status (checked on 3 October 2026).

This general information does not replace the official texts: check the regulations in force before buying or exchanging plants or seeds, ask the seller for proof of origin and prefer plants raised from cultivated seed. Vannini (2019) notes that both wild-collected and artificially propagated seed have been easy to obtain in the United States.

Growing Zamia nesophila

Zamia nesophila is one of the more forgiving plicate-leaved Zamia, and one of the few for which growers have published results, almost all on young plants. Simon Lavaud, of the French specialist nursery Cycadales.eu, places it in his group of lowland Zamia of Central and South America, which he rates difficult outside the tropics, but singles it out, with Zamia obliqua, as a tree-forming species regularly offered by nurseries and fairly easy to grow. For that group he recommends free-draining compost, neutral to acid water, nights at about 22 °C (71.6 °F), with brief drops to about 15 °C (59 °F) tolerated, days at 28 to 30 °C (82.4 to 86 °F) and high air humidity, ideally 70 to 90% (Lavaud, 2023).

The advice given on the Fous de palmiers forum points the same way:

  • Warmth: the grower who distributed the seeds advises reducing watering, without letting the compost dry out, as soon as the temperature drops below 20 °C (68 °F); in a conservatory kept at 21 °C (69.8 °F) all year, his plants grew well (2013).
  • Humidity: the more humid the air and the deeper the shade, the broader the leaflets; good air movement is considered essential, as stagnant air encourages rot.
  • Light: partial shade. The species is said to take a little more sun than Zamia hamannii, but its leaves yellow in full sun. A grower in Saône-et-Loire, in eastern France, reports on the Fous de palmiers forum that in July 2015 his young potted plants, placed in partial shade, had scorched leaves, and that they recovered in full shade; another, in Finistère (Brittany), saw weaker growth and paler leaves on a plant moved into a glasshouse in summer 2015, which he put down to the heat.
  • Compost: very free-draining and mostly mineral, sandy as in the habitat. A grower in the Dordogne lost three plants out of five in 2015, their roots eaten by beetle larvae in a very organic potting compost.
  • Feeding: forum growers recommend a slow-release fertiliser rich in nitrogen.

Outside Europe, a grower in French Guiana judged in 2014 that Zamia nesophila was “the fastest of all” the species he had sown. In Florida, Vannini (2019) reports that it apparently performs well in the landscape in the south of the state, showing promise as a garden plant and tolerating more direct sun than its close relatives from western Panama. Jungle Music, a Californian nursery, recommends filtered light rather than direct sun and notes that the plant grows in sandy soil in the wild.

Outdoor and in-ground cultivation

Outdoors it is a plant for frost-free tropical and humid subtropical gardens: southern Florida, Hawaii, northern Queensland. Its salt tolerance in the wild makes it a candidate for coastal plantings, though no trial in saline soil has been published. Give it free-draining, sandy soil, dappled shade at first and room for a crown of leaves well over 2 m long.

Container cultivation

Elsewhere it is grown in a warm glasshouse or conservatory, which suits it well while young. Use a deep pot of very free-draining, mostly mineral compost that is never allowed to dry out completely, keep it warm by night as well as by day, in partial shade with humid but moving air, and reduce watering without stopping it when temperatures fall. In time the size of the leaves becomes the main constraint.

In cultivation in the UK, US and Australia

In short: widely held in American gardens, present in Sydney; no British record.

  • United States: Montgomery Botanical Center (Coral Gables, Florida) lists 24 accession numbers under Zamia nesophila in its online catalogue, from 2001 to 2008, plus seven under Zamia aff. nesophila (queried on 25 October 2026); Fairchild Tropical Botanic Garden (Coral Gables) lists it in its online Garden Explorer. BGCI also records it at Marie Selby Botanical Gardens (Sarasota), Naples Botanical Garden and Edison and Ford Winter Estates (Fort Myers), in Florida; Atlanta Botanical Garden, Georgia; San Diego Botanic Garden (Encinitas, California); Honolulu Botanical Gardens, the University of Hawaii at Hilo and the National Tropical Botanical Garden (Kalāheo), in Hawaii; and the conservatory of the University of Minnesota, St Paul (queried on 25 October 2026).
  • United Kingdom: no record in BGCI or in the online catalogues of British gardens.
  • Australia: the Royal Botanic Garden Sydney lists it in its online catalogue (Garden Explorer, updated on 1 October 2026) and declares it to BGCI.

A BGCI declaration shows that a garden held the plant at some point, not that it is alive there today, and these sources do not say whether the plants are grown outdoors or under glass. In Europe, Prague Botanical Garden also declares it.

Propagation

The plant is dioecious: a male and a female plant are needed to obtain seed. Fresh seed germinates quickly and easily: members of the Fous de palmiers forum obtained germination in one to two weeks, sometimes three days, at 25 to 28 °C (77 to 82.4 °F), in a bag of moist sphagnum, peat, vermiculite or coir. Reported rates range from 4 out of 12 (a batch of seeds without embryos, 2010) to 9 out of 10 (2015). The grower who distributed them cleaned off the fleshy coat with an enzyme, pectinase, and advises burying the seed halfway, germination end down, with the apex about 1 cm below the surface so that it does not dry out.

The original description also records how the trunks regrow when cut: severed tops re-root and continue to grow, and stumps produce new buds (Taylor and colleagues, 2008), but no propagation by cuttings has been documented in cultivation. Seeds are exempt from CITES under annotation #4; make sure they come from cultivated plants.

Pests and diseases

In cultivation, forum growers report seed or apex rot just after germination, especially when the heat is excessive or irregular, and beetle larvae in composts rich in organic matter. In the wild, caterpillars of the hairstreak butterfly Eumaeus godarti feed on new and old leaves in two populations, apparently without serious harm, and a fungal leaf spot (possibly Mycoleptodiscus indicus) affects a few plants growing in the open, in sunnier spots (Taylor and colleagues, 2008).

Cold hardiness

Zamia nesophila is not hardy: it is a tropical beach plant from a place where the temperature rarely falls below 18 °C (64.4 °F). Two reports nevertheless show that young plants, or plants sheltered by a tree canopy, can survive light frost at the cost of their leaves.

Mediterranean and European experience

A grower on the Côte Bleue, the coast west of Marseille (Bouches-du-Rhône, France), reports on the Fous de palmiers forum that his young plants of Zamia nesophila, sown at the beginning of 2013 and planted out in shade under tropical foliage in summer 2014, “seemed to be getting through” the winter in January 2015, then, in September 2015, that they had come through the winter of 2014–2015 “without trouble”. He gives no temperature: the nearest Météo-France station, Marignane, 10 to 20 km away depending on the village, recorded a minimum of −3.0 °C (26.6 °F) on 10 February 2015; it is not on the coast, and the temperature the plants actually experienced is unknown. The grower who had supplied the seeds called this planting “daring”: a defoliated plant survives but loses time.

In February 2015 the same seed distributor reported on the forum that a correspondent in Greece grew Zamia nesophila in the ground under a tree canopy, in an area with minimums of −2 to −2.5 °C (28.4 to 27.5 °F): the plants lose their leaves in winter and regrow in spring, but grow slowly. This is a second-hand report, with no precise place or dated episode. All the other French results are for potted plants brought indoors for winter; a grower in Loire-Atlantique took his in on 13 October 2013, when the temperature fell to 6 °C (42.8 °F).

For readers in Mediterranean-type climates such as coastal California or southern Australia, these results suggest that a young plant in a sheltered, shaded spot may survive a brief light frost, but it will be set back; a frost-free glasshouse remains the safe choice.

United Kingdom, United States and Australia

No cold record has been published from these countries. The American gardens that hold it are mostly in southern Florida, coastal southern California and Hawaii, and in Australia in Sydney; none says whether its plants grow outdoors or under glass. In the UK it needs a heated glasshouse all year.

Toxicity and precautions

No toxicity study specific to Zamia nesophila has been published. The risk is, however, well documented in cycads: Zamia contain toxic glycosides (cycasin, macrozamin), from which methylazoxymethanol is derived, a compound toxic to the liver and nervous system, and the ASPCA cites cycasin and BMAA, a neurotoxic amino acid, for Zamia furfuracea, which it lists as toxic to dogs, cats and horses; the seeds are the most dangerous part. The local use of the trunk as a drink (see below) does not make the plant safe. In the absence of evidence to the contrary, treat the plant as poisonous: keep seeds and leaves out of reach of children and animals, and wear gloves to clean seeds. If any part is swallowed, contact a poison centre, a doctor or a vet straight away.

Traditional uses

On the islands, the trunks are cut, ground up and brewed into a tea that is drunk as a sexual stimulant. According to accounts gathered on two islands, inhabitants of African descent said that only men use it, to prolong erections, while the indigenous inhabitants of a nearby island said that both men and women enjoy its effects (Holzman and Haynes, 2004, cited by Taylor and colleagues, 2008). The authors stress that all cycads are highly toxic and that the long-term effects of this use are unknown. Repeated cutting of the trunks explains the true “lawn” of young plants found near houses: severed tops that fall to the ground take root, and the stumps form new buds, giving multi-headed plants.

Frequently asked questions

Where does Zamia nesophila grow?

Only on a few islands of the Bocas del Toro archipelago, in north-western Panama, mostly in beach sand just above high water, and less often on low forested hills.

Does it tolerate salt?

Yes, in its habitat: the largest populations receive salt spray and are sometimes flooded by seawater. Prolonged submersion does rot the roots, according to an amateur who visited the habitat, and no trial in saline soil has been published.

Is it hardy?

No. Young plants survived the winter of 2014–2015 in the ground, in shade, near Marseille, where the nearest station recorded −3.0 °C (26.6 °F), but this is an isolated result. Treat it as a frost-free plant, or a glasshouse plant in the UK.

How do I tell it from Zamia hamannii?

By the colour of the new leaves: bright green and hairless in Zamia nesophila, pinkish brown to pinkish orange and silvery in Zamia hamannii. Both live on the islands of Bocas del Toro.

Are the seeds easy to germinate?

Yes, if fresh: forum growers obtained germination within one to two weeks at 25 to 28 °C (77 to 82.4 °F), in a bag of moist sphagnum or vermiculite.

Sun or shade?

Partial shade, at least for young plants. It takes more sun than its close relatives, and plants in southern Florida tolerate direct sun, but seedlings raised in shade scorch quickly in direct sun.

Is it endangered?

Yes. The IUCN rates it Critically Endangered (2022): populations of several thousand plants have been destroyed by coastal development.

Is it poisonous?

It must be assumed so. No analysis specific to the species exists, but the cycads studied contain serious toxins, mainly in the seeds.

Further reading on Succulentes

Sources

Reference sites

Bibliography

  • Taylor B., A.S., Haynes, J.L. & Holzman, G. (2008). Taxonomical, nomenclatural and biogeographical revelations in the Zamia skinneri complex of Central America (Cycadales: Zamiaceae). Botanical Journal of the Linnean Society 158(3): 399–429. https://doi.org/10.1111/j.1095-8339.2008.00886.x [Original description; full text read.]
  • Taylor B., A.S., Haynes, J.L., Stevenson, D.W., Holzman, G. & Mendieta, J. (2012). Biogeographic insights in Central American cycad biology. In: Stevens, L. (ed.), Global Advances in Biogeography, InTech, Rijeka: 73–98. [Chapter read in full.]
  • Tang, W., Skelley, P.E., Taylor B., A.S. & Salzman, S. (2024). Review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting cones of the cycad Zamia L. (Cycadales) in Panama, with descriptions of five new species. Insecta Mundi 1043: 1–40. [Cone beetles; full text read.]
  • Skelley, P.E., Tang, W., Salzman, S. & Calonje, M. (2026). Taxonomic review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting cones of the cycads Zamia L. and Microcycas (Miq.) A.DC. (Cycadales) in the Caribbean Basin and Central America north of Panama, with descriptions of five new species. Insecta Mundi 1184: 1–48. https://doi.org/10.64338/im.1184.9bdte [Confirms Pharaxonotha clarkorum as the sole known beetle pollinator; full text read.]
  • Lindstrom, A., Habib, S., Dong, S., Gong, Y., Liu, J., Calonje, M., Stevenson, D. & Zhang, S. (2024). Transcriptome sequencing data provide a solid base to understand the phylogenetic relationships, biogeography and reticulated evolution of the genus Zamia L. (Cycadales: Zamiaceae). Annals of Botany 134(5): 747–768. https://doi.org/10.1093/aob/mcae065 [Full text read on PubMed Central.]
  • Taylor-Blake, A., Haynes, J. & Holzman, G. (2022). Zamia nesophila. The IUCN Red List of Threatened Species 2022: e.T187842A1829425. https://doi.org/10.2305/IUCN.UK.2022-1.RLTS.T187842A1829425.en [Red List assessment; reference given by The World List of Cycads and checked on Crossref; the Red List page could not be consulted.]
  • Vannini, J. (2019). Central American and Panamanian cycads in field and greenhouse. Exotica Esoterica, 17 February 2019 (updated 2026). [Naturalist’s article; read.]
  • Lavaud, S. (2023). Comment cultiver les Zamia en climat tempéré. Cycadales.eu. [Growing advice for groups of species; read directly.]
  • Haynes, J.L. (2022). Etymological compendium of cycad names. Phytotaxa 550(1): 1–31. https://doi.org/10.11646/phytotaxa.550.1.1 [Source of the etymology given by The World List of Cycads; not consulted directly.]
  • Calonje, M., Stevenson, D.W. & Osborne, R. (2026). The World List of Cycads, version 2026.08.15. Montgomery Botanical Center, Coral Gables. https://doi.org/10.5281/zenodo.21958524 [Reference checklist; consulted directly.]