Cycads are an ancient group of Gymnosperms, often considered to be living fossils. Cycads originated over 290 million years ago and related to the extinct Cycadofilicales (seed ferns) from which they evolved. The Order Cycadales currently includes three families with living members: Cycadaceae, Stangeriaceae and Zamiaceae.
While Cycads aren't succulent they are often grown by collectors of cacti and other succulent plants. If the smaller species are selected it is possible to grow a mature plant of a reasonable size for a small glasshouse. Cycads should be generally regarded as poisonous to livestock as they contain the unique methylazoxymethanol-glycoside neurotoxins Cycasin and Macrozamin. There does not seem to be a problem with growing and handling Cycads as long as no parts are eaten, although it might be wise to use gloves when cleaning the seeds as these contain the highest concentration of toxins.
Despite their toxicity, Cycad nuts, seeds or the starchy pith of some species have been used as traditional foods after thorough washing or other treatment to remove neurotoxins. Guam Dementia is a late onset neurodegenerative disease thought to be caused by consumption of flour made from improperly washed starch extracted from cycad seeds, during and after World War II when there were food shortages.
A large collection of Cycads, including the National Collection of Encephalartos, can be seen at the Royal Botanic Gardens, Kew, especially in the Palm and Temperate Houses.
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Cycadaceae Persoon 1807
This family of Cycads appeared in the Cenozoic era over 65 million years ago, possibly diverging from the other Cycads of the Family Zamiaceae in the earlier
Mesozoic. The Madagascar Sago Palm Cycas thouarsii is thought to be the oldest living Cycad, with fossils dated as 140 million years old.
The type genus Cycas is the only genus currently recognised in the family Cycadaceae and is only distantly related to members of the other Cycad families.
Cycadaceae usually have thick, pithy stems that rarely branch. However, they may offset at their base. Much of the structural support is provided by persistent woody leaf bases around the stem. The alternate pinnately-compound leaves are clustered in a spiral at the apex of the stem, each having many stiff, waxy leaflets either side of the leaf mid-rib. Leaves emerge from the apical bud with leaflets tightly coiled inwards like a watch spring. Each leaflet has a single mid-vein.
Cycads produce male cones and female ovules on separate plants (dioecious). Male plants produce cones with many scales, each of which has microsporangia (pollen) on its lower surface. A distinctive feature of the Cycadaceae is that female plants don't form cones like other cycads. Instead they produce ovules on the lower surface of modified leaves arranged in two to three whorls at the stem apex, with up to four female megaspores in each ovule. Seeds are usually large.
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Cycas Linnaeus 1753
Name: Greek Koikas = a kind of palm (Theophratus)

The type genus Cycas includes about 95 species of woody perennials with a palm-like appearance. Cycads have a cylindrical caudex or trunk, sometimes almost entirely subterranean, with a spiky appearance from persistent petiole bases on its surface and with a crown of thick, hard pinnate leaves arranged spirally at its end. Young parts of the plant are hairy. The leaflets are distinctive, with a prominent midrib and without any obvious secondary veins. When leaves emerge, the leaflets are coiled like watch springs.
Cycas are native to tropical and subtropical regions of the Old World. They are found in eastern and southeastern Asia including the Philippines (10 species), Eastern Africa including Madagascar, Northern Australia, Polynesia, and Micronesia, Australia (26 species), while the Indo-Chinese area (30 species).
Cycads grow slowly, becoming mature and producing cones only after at least 10 years growth. Most species are tropical and require heat to grow well. However the Sago Palm Cycas revoluta is undemanding and flourishes in a cool conservatory.
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Cycas revoluta Thunburg 1782 (King Sago Palm)
Name: leaflets curl under along their edge = revolute

This is probably the most widely-cultivated Cycad. It's attractive feathery foliage adds an exotic touch to any conservatory. Even a small plant has a conical caudex which, with age slowly elongates into an unbranching trunk, and may offset at it's base to form a clump. A several centuries old plant may develop a 6 ft trunk.
Cycas revoluta is native to the southern Japan islands of Kyushu and Ryukyu, growing further North than any other species of Cycad. It is a very hardy plant, tolerating drought, and light frosts and is almost hardy in South-East England through an average Winter. Several specimens survive planted in sunny corners outside the Temperate House in Kew Gardens. A plant should not be moved while it's leaves are unfurling as they will become distorted, and being long-lasting, will spoil the plant's very symmetrical appearance for some years.
All parts of the Sago Palm Cycad are extremely poisonous to animals if ingested. The seeds contain the highest level of cycasin and other toxic metabolites.
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 Sago Palm Cycas revoluta

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Cycas siamensis Miquel 1863
Name: Siam is an old name for Thailand
This Cycad produces flat leaves whose colour (silvery-white, yellow, orange, brown) depends on the colour of hairs on the leaves.
Native to Thailand and Vietnam. All parts of the plant are poisonous if ingested
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Cycas thouarsii R. Brown ex Gaudichaud 1829 (Madagascar Sago)
Named for: French botanist Louis-Marie Aubert du Petit-Thouars (1758-1831)

This Cycad has a stout trunk over 30ft tall topped with a crown of up to 40 feathery pinnate leaves. Offsets may be produced along the length of the trunk.
Native to coastal Kenya, Tanzania and Mozambique, Madagascar, Comoros Islands and introduced into the Seychelles. This Cycad probably originated over 140 million years ago and may be the most ancient living species. It is relatively fast-growing, tolerating light frosts.
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Boweniaceae Stevenson 1981
a small family of Cycads in which the genus Bowenia (2 living species) is sometimes placed. The Boweniaceae are distinguished by the unique branching midrib (rachis) of their leaves.
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Bowenia Hooker ex Hooker fils. 1863
Named for: Sir George F. Bowen 1821-1899 first governor of Queensland.

The genus Bowenia includes two living species of small fern-like shrubs from Eastern Queensland, Australia. This genus is variously included in its own family Boweniaceae or with the Stangeriaceae or Zamiaceae. The leaves are bipinnate with leaflets lacking a midrib and not articulated. Two additional species in the fossil record are known from Australian coal deposits.
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Bowenia serrulata Chamberlain 1912 (Byfield Fern) Syn. B. spectabilis Bull 1878
Name: Latin serrulatus = serrated or toothed, referring to the leaf margins.

The 10 - 12 in globular subterranean stem produces up to 20 branching bipinnate leaves 4 - 5 ft tall, whose leaflets with finely-serrated edges. The stem has a large tap root.
Native to warm, dry Eucalyptus forests of Eastern Queensland where it grows as an understorey plant.
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Bowenia spectabilis Hooker ex Hooker fils. 1863 (Zamia Fern)
Name: Latin spectabilis = remarkable, showy

The type species for the genus, native to subtropical moist forests in North-Eastern Queensland. The underground stem produces up to seven 4 - 5 ft branching, bipinnate leaves whose leaflets have smooth margins.
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Stangeriaceae Johnson 1959
a small family of Cycads with the genus Stangeria (1 living species) and sometimes including the genus Bowenia (2 living species). Distinctive features of the family are vascularised stipules and almost complete absence of cataphylls. Bowenia may be destined for its own separate family Boweniaceae.
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Stangeria Moore 1853
Named for: Dr Max Stanger, Surveyor General of Natal province, South Africa, who collected a specimen in 1851.

Stangeria is monotypic for the fern-like cycad Stangeria eriopus, although the fossil record suggests that this genus was once distributed globally.
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   Stangeria eriopus soft-leaf forest form |
Stangeria eriopus Baillon 1892 (Natal Grass Cycad)
Name: Greek erio = woolly + pus = footed, referring to the woolly petiole base.

The thick stem is subterranean, with a tap-root and corralloid roots near the surface housing nitrogen-fixing cyanobacteria. The stem is crowned with distinctive pinnately-veined, fern-like leaves. Indeed this Cycad was originally described as a fern (Lomaria eriopus Kunze 1839) and the mistake only realised when seed cones were observed. This variable Cycad has a forest form (above) bearing soft, thin, wide 6ft leaves and a more compact form (below) from open grassland with smaller, thicker 1ft long stiff leaves.
Stangeria eriopus matures relatively quickly for a Cycad, usually after 5 - 7 years, from which time male and female stalked cones are produced on separate plants. The cones produce a perfume to attract beetle pollinators.
Native to coastal Eastern South Africa and Southern Mozambique, although not within range of salt spray. Stangeria eriopus, especially the root, is a traditional medicine as a purgative and for treating headaches
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     Stangeria eriopus hard-leaf form. Left: grown in shade. Centre, Right: grown in strong light. |
Encephalartos Lehmann 1834 (Bread Palms)
Name: Greek words en = within + kephali = head + artos = bread

The genus Encephalartos includes 68 species of African Cycads, primitive Gymnosperms. The majority of species are in Southern Africa with 37 species found in South Africa. A few species are found in Angola, Benin and Ghana. All species of Encephalartos are endangered.
The starchy pith in the centre of the stem of several species of Encephalartos can be used to prepare a native bread, hence the common name. Toxins in the pith are destroyed by prolonged burial in the ground.
The Royal Botanic Gardens, Kew houses the National (UK) Collection of Encephalartos.
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Encephalartos altensteinii Lehmann 1834 (Eastern Cape Giant Cycad, Bread Tree )
Named for: Baron von Stein zum Altenstein, Prussian statesman.

This large impressive Cycad has a single erect trunk up to 15ft tall, often suckering at the base to form a clump with age. The pinnate 10ft long, deep green glossy leaves are long-lived, but a new flush of leaves is produced in the Spring. The broad, well-spaced leaflets have 1-3 prickles on their margin, but basal leaflets do not merge into spines leaving a short length of bare petiole. The midrib of the leaf is straight, not twisted.There is no wooly fibre in the crown around new leaves. Yellowish male and female cones are produced at the stem tips of separate plants.
Encephalartos altensteinii is native to the South-Eastern Cape of South Africa, a Summer rainfall area and a guide to watering the plant. It is commonly grown in tropical gardens, but is very slow-growing. It will not stand more than the lightest frost.
Left: This huge multi-stemmed E. altensteinii in the Kew Palm House arrived at Kew in 1775 and is claimed as the oldest pot plant in the world, although by no means the oldest specimen in cultivation. A specimen of E. altensteinii in Lednice Greenhouse, Czech Republic is 400 - 500 years old.
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Encephalartos arenarius Dyer 1955 (Alexandria Cycad)
Name: Latin arenarius = growing in sandy places

Encephalartos arenarius has a stout 4 ft stem that may be either subterranean in leaf litter or prostrate. The plant clumps up by offsetting at the base. The 4 ft pinnate leaves are recurved at their ends and consists of many broad, hard leaflets with prickly lobes. Two forms of E. arenarius are known with either dull green or bluish leaves.

Native to coastal sandy dune forest around Alexandria in the Eastern Cape of South Africa, although much habitat has been lost from conversion to pastures.
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Encephalartos cupidus Dyer 1971
Name: Latin cupidus = bristly

The mainly subterranean main stem produces 3ft pinnate blue-green leaves. The leaflets form a V-shape on the mid-rib and are reduced to vestigal prickles on the petiole. They have 4 - 6 spines on each margin. The cones are greenish.

Native to the Eastern Transvaal Drakensberg escarpment, occurring as small populations. Plants may clump up by suckering.
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Encephalartos ferox Bertolini 1851 Syn. E. kosiensis (Zululand or Tongaland Cycad, Tongaland Broodboom)
Name: Latin ferox = ferocious, referring to the rigid, spiny leaflets.

This medium sized Cycad has a stout 3 - 4 ft long trunk which can be mostly subterranean, pulled down by contractile roots.
The pinnate leaves have hard ovate blue-green leaflets with spine-tipped lobes, forming a shallow V-shape along the mid-rib. The male and female cones on separate plants are a dark salmon-pink.

Encephalartos ferox is native to coastal Mozambique and Natal where it grows on sand dunes and evergreen forests. The starchy pith has been used to make a flour after fermentation to remove toxins.
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Encephalartos horridus Lehmann 1834 (Eastern Cape Blue Cycad)
Name: Latin horridus = bristly

This low-growing Cycad has the majority of its large 3ft long stem underground, although more may be exposed when growing in rocky terrain. Two forms are recognised: a robust form and a dwarf form with shorter stems and smaller leaves. The freely-branching stem is crowned with 4 - 8 stems of distinctive blue-grey leaves with stiff, spiny leaflets. After becoming mature, each crown of the plant produces a single brown or dark red cone covered in fine hair.

Female cones are egg-shaped and male cones cylindrical with narrow ends.
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Female cones produce toxic seeds surrounded by a fleshy skin which is eaten by birds and monkeys, while discarding and dispersing the more toxic seeds.

Native to the Eastern Cape of South Africa around Port Elizabeth and Uitenhage, growing on acidic soils. Because of its unique appearance, natural populations have been depleted by over-collection. Not cold-hardy but may tolerate brief, light frosts.
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Encephalartos lanatus Stapf & Burtt Davy 1926 (Woolly Cycad, Olifants River Cycad)
Name: Latin lanatus = woolly

This medium-sized Cycad has a globular caudex becoming a thick 4ft tall stem with great age. The stem bears many leaf scars and is crowned with blue-grey 3ft leaves which are covered in silver hairs when young. The leaflets for a V-shape along the mid-rib and the leaves tend to recurve. The cones are also covered with persistent silvery hairs.

Native to limited locations in the Transvaal at altitudes of 4000 - 5000 ft. This slow-growing Cycad is frost hardy and adapted to both drought and bush fires but dislikes too much water.
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Encephalartos lehmannii Lehmann 1834 (Karoo Cycad)
Named for: Professor J.G.C. Lehmann of Hamburg who created the genus.

This Cycad has a stout trunk up to 6ft tall, crowned with a whorl of beautiful feathery blue leaves 3 - 4 ft long. The stems clump up from the base.

Native to the Eastern Cape Province of South Africa. One of the most drought resistant Cycads and tolerates a degree of frost.
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Encephalartos longifolius Lehmann 1834 (Thunberg's Cycad, Suurberg Cycad, Bread Palm, Suurbergbroodboom)
Name: Latin longis = long + folius = leaf

The thick trunk up to 15ft tall, produces a crown of glossy, dark-green leaves curving outwards towards their ends. The lanceolate leaflets are arranged on the mid-rib in a distinctive V-shape. Their lower margin may have a small number of teeth. This species clumps up by offsetting from the base. The large olive-green female cones are the heaviest of any Cycad. Male cones are brownish. The seeds are very toxic.

Native to the Eastern Cape. The starchy pith was fermented to remove toxins and used to make a flour. One of the first Cycads to be discovered by the Swedish botanist Carl Peter Thunberg and Francis Masson in 1772.
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Encephalartos natalensis Dyer & Verdoorn 1951 (Natal Giant Cycad, Natalbroodboom)

The stout trunk can grow 20ft tall and is crowned with 10 ft long pinnate leaves with many well-spaced bright-green ovate leaflets, usually but not always without marginal prickles. Basal leaflets are reduced to spines along the petiole. New leaves and cones are covered in yellowish hairs. The cones are yellow and ovoid. Female cones contain toxic red seeds.

Native to the Eastern Cape Province and KwaZulu-Natal. The starchy pith of the trunk was processed into a flour. This Cycad is closely related to E. altensteinii.
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Macrozamia Miquel 1842 (Burrawang)
Name: Greek macros = "large" plus the related genus Zamia

Most of the approximately 40 species in the genus Macrozamia occur in Eastern Australia in New South Wales and Queensland but one species is native to the Northern Territory and three species are from Western Australia. All species have dark green, palmate fronds arising from a central trunk. In some species the trunk is very short or subterranean. Macrozamia is distinguised by pinnate leaves in which the leaflets have numerous parallel veins but lack a midrib and have a coloured swollen base. Lower leaflets may be present as spines. Leaves are interspersed with cataphylls and are pubescent especially when young. The sporophylls are arranged spirally in the male cones. The female cones have megasporophylls terminating with an upward spine.
Despite their toxicity to humans and livestock, several species of Macrozamia were used as food plants by Aborigines. The starchy seeds are extremely poisonous and considerable masceration in water over a period of a week, with several changes of water, was needed to remove the toxins. The resulting pulp was used as flour to make cakes. The leached toxins were used as fish poisons.
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Macrozamia communis L.A.S.Johnson, 1959 (Burrawang)
Named for: habit of growing in large, dense groups

The caudex is mostly subterranean but can extend above ground as a short trunk in old plants or where the substrate is impenetrable.
Native to the open forests along the Eastern coast of New South Wales, extending inland to the slopes of the Great Dividing Range. The common name 'Burrawang' really belongs to this plant although it is often applied to other species of Macrozamia. The toxic yellow to red seeds were processed into flour by the Aborigines.
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Macrozamia miquelii de Candolle 1868
Syn. Macrozamia miquelii Mueller 1862
Named for: F.A.W. Miquel, Dutch botanist and physician

The caudex is mostly subterranean but can extend 2 - 3 ft above ground in old plants.
Native to coastal Southern Queensland and Northern New South Wales.
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Macrozamia moorei von Mueller 1881 (Moore's Cycad, Bottle Tree)
Named for: Charles Moore (1820-1905), director of the Royal Botanic Gardens, Sydney.

Macrozamia moorei is the tallest Macrozamia with a massive 3ft diameter trunk up to 20 ft tall and covered with persistent leaf bases. The arching pinnate leaves up to 8ft long have numerous narrow leaflets arranged in a V-shape along the midrib. The lower leaflets degenerate into spines along the short petioles. Male and female cones are on separate plants. The barrel-shaped female cones contain many large scarlet seeds, which are extremely poisonous. However, Aboriginals made flour from the seeds after lengthy soaking in water to remove the toxins.
Native to central Queensland. This very slow-growing Cycad is propagated as a conservatory plant or for landscaping in Mediterranean climates.
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Zamia Linnaeus 1763
Name: Greek azaniae = "pine cone" Latin zamia = "pine nut"

The genus Zamia includes about 50 species of small, shrubby Cycads, with a palmate appearance and stout stems which may be subterranean or up to 20ft tall. The stems are crowned with spirally-arranged pinnate leaves with a degree of pubescence on younger leaves. The broad leaflets lack a midrib, and have subparallel dichotomous venation. Petioles may have spines. Zamia picta is noteworthy for variegated leaves with pale speckles. Zamia pseudoparasitica grows as an epiphyte on trees.
Mature plants produce cones with vertical rows of sporophylls. Seeds are fleshy and globular to oblong in red, orange to pale yellow.
Members of the genus Zamia are native to a wide range of both North, and South America with most species coming from Central America. Zamia integrifolia grows as far North as Georgia and Zamia boliviana as far South as Bolivia. Most species grow as understorey plants in warm humid rain forests and are not frost hardy.
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Zamia furfuracea Cardboard Palm
 Cardboard Palm Seeds
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Zamia furfuracea Linnaeus fils. 1789 (Cardboard Palm) Syn. Z. maritima
Name: Latin furfuracea = "mealy, scurfy"

The short, partly subterranean trunk is crowned with stiff pinnate leaves composed of up to twelve stiff, pubescent leaflets, sometimes with toothed tips.
Mature plants produce redish-brown cones in the centre of the plant. Pollination is by the belid weevil Rhopalotria mollis. Female cones produce fleshy, crimson-coloured seeds which are very poisonous if ingested. |
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Native to the Mexican state of Veracruz, growing in sand dunes where it tolerates some salt spray. A slow growing plant when young but tolerant of cultural conditions. May tolerate very light frosts, with regrowth of damaged leaves. This species is widely grown as a houseplant.
All parts of the plant are poisonous if ingested, causing liver and kidney failure and paralysis. There is no antidote.
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 Z. integrifolia |
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Zamia integrifolia Linnaeus fils. 1789 (Coontie, Florida arrowroot, wild sago) Syn. Z. floridana, Z. pumila

Name: Latin integrifolia = entire + folium = leaf, refers to leaf margins without teeth (not quite correct). Seminole Native American conti hateka = "white flour root" -> Coontie (or koonti).

Zamia integrifolia has a short 10 in trunk, often mostly subterranean, and a substantial tuberous root. The stem is crowned with pinnate leaves formed from up to thirty pairs of lanceolate leaflets which may have indistinctly toothed tips. Petioles are often prickly and young leaves are covered in reddish-brown hairs. In a temperate climate the leaves can be deciduous during cold periods, making this species relatively hardy to light frosts. Separate plants produce clusters of cylindrical male cones and ovoid female cones.
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     Z. floridana Z. pumila |
Zamia integrifolia is the only Cycad native to the South-Eastern USA, growing on limestone or sandy soils as far North as Florida and Georgia. Also native to the Bahamas, Cuba, Grand Cayman, Hispaniola, and Puerto Rico although populations have been depleted in parts of the original distribution. Some authors split this variable species into others e.g. Zamia floridana, Z. augustifolia, Z. floridana, Z. pumila, Z. silvicola, and Z. umbrosa.

All parts are poisonous if ingested, especially the fleshy red seeds. However, the common name refers to use of the Coontie, with considerable processing, to make an edible starch. Zamia integrifolia and related Zamias are the sole food plants for caterpillars of the endangered butterfly Eumaeus atala.
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Zamia loddigesii Miquel 1843 Syn. Zamia cycadifolia Dyer 1884
Named for: Conrad Loddiges, English nurseryman.

This Cycad has a small, usually subterranean caudex bearing up to 6 pinnate leaves supported on spiny petioles. The stiff leaflets are lanceolate with slightly toothed margins.
Native to sandy soils at modest elevations in Eastern Mexico and Guatemala, where it is common but threatened by habitat loss. A drought tolerant plant suitable for xerophytic landscaping.
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Zamia muricata Willdenow 1806 Syn. Z. verschaffeltii, Z. monticola

The short stem, branching with age, produces a crown of up to six upright leaves which increase in length and number of leaflets as the plant matures. This is a relatively fast-growing plant, becoming mature with production of cones after 5 - 6 years.

Native to low-level rainforests of Colombia and Venezuela where it grows as an understorey plant in wet clay soils.
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Zamia neurophyllidia Stevenson 1993
Name: refers to the nerve-like network of grooves in the leaves

This small plant has a short trunk producing up to ten pinnate leaves with small prickles along their margins and prickles along the petiole. Very similar to Z. skinneri from which it was separated, but is smaller and with fewer chromosomes.
Native to low-altitude tropical rainforests of Costa Rica, Nicaragua and Panama. Needs moist but not soggy soil.
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Zamia vasquuezii Stevenson, Sabato, Moretti & De Luca 1995 (Chamalillo, Teocinte)
Named for: Mario Vazquez Torres, Mexican botanist who discovered this species.

A small Cycad with a subterranean trunk producing a crown of 3 ft pinnate leaves with more than 12 pairs of leaflets with toothed margins and small spines on the petioles. Similar to the smaller Z. fischeri, from which it was segregated. Z. fischeri has fewer, smaller leaves with less than 12 pairs of leaflets.
Native to pine - oak forests in the Northern part of the Mexican state of Veracruz although severely depleted in its habitat.
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