Dioon

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Dioon
Dioon edule01.jpg
Dioon edule
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Cycadophyta
Class: Cycadopsida
Order: Cycadales
Family: Zamiaceae
Subfamily: Encephalartoideae
Tribe: Diooeae
Schuster
Genus: Dioon
Lindl. [1] [2]
Type species
Dioon edule [1]
Lindl.
Dioon Distribution.png
Synonyms [3]

Dioon is a genus of cycads in the family Zamiaceae. It is native to Mexico and Central America. [3] Their habitats include tropical forests, pine-oak forest, and dry hillsides, canyons and coastal dunes. [4] [5] [6] [7]

Contents

Description

Dioons are dioecious, palmlike shrubs with cylindrical stems, usually with many leaves. The species in the genus Dioon are perennial, evergreen cycads with cylindrical stem axis is partially in the ground. The plant is thickened and made of soft wood, rarely having above ground branches. Leaf bases are persistent or shedding to leave smooth bark. The leaves are pinnate, spirally arranged, interspersed with cataphylls, with leaflets not articulated and lacking a midrib. The lower leaflets are often reduced to spines. The sporophylls are not in vertical rows in cones, and the megasporophyll apices are broadly flattened, upturned, and overlapping.

Species in the genus Dioon have 2n = 18 chromosomes. [8]

The largest species is D. spinolosum , which are over 16 meters high and whose trunk diameter can reach 40 centimeters. However, typical specimens of other species are only between three and six feet high or remain even smaller. Dioon can get very old, maybe even over 1000 years. Dioon edule and Dioon tomasellii have the most widespread ranges. Most species have highly limited geographical ranges.

Leaves

Leaf of Dioon spinulosum with are identified by the veining and prickly margin Dioon spinolosum04.jpg
Leaf of Dioon spinulosum with are identified by the veining and prickly margin

The leaves are paired pinnate and are spirally on the stem axis. Some leaves are initially wrapped in protective leaves during their development, which are called cataphyll. Unlike other cycads such as Stangeria eriopus , the juvenile fins are not curled either transversely or longitudinally, but straight. The lower leaflets are often reduced to their petiole. The petioles are uncorned and thickened at the base.

The leaflets are simple, often with a prickly edge. The primary nerve consists of many forked dividing nerves, without a recognizable midrib. The nerves spring directly from the edge of the leaf rhachis and then run towards the plumage axis. The spars are hairy at least on young leaves. The hairs (trichomes) are colorless, branched or simple.

The stomata for the gas exchange are found either only on the underside of the leaf, or in some species on both sides. Glands are not otherwise colored and difficult to identify. The cells of the leaf epidermis are extended parallel to the plumage axis.

Roots

Like other cycads, coral-like roots are sometimes formed that grow up from the primary root and branch out quickly just below the soil surface. The individual roots are thickened rhizomes and are inhabited by cyanobacteria of the genus Nostoc, which mainly used them for nitrogen fixation. There also seems to be a symbiotic relationship with arbuscular mycorrhizal fungi. [9]

Cones and Seeds

All Dioons are dioecious, having either male or female organs, but only on different individuals. The male cone is stalked. The male cone scales called microsporophyll, which are flattened and bent upwards, sterile at the tip . They are arranged spirally around the pin axis. Each microsporophyll carries on its underside (abaxial side) a variety of pollen sacs. These open with slots, from which the pollen then escapes. The pollen grains are streamlined and have only one germ line (monosulcate). The male cones fall off after one year.

The female cones are also stalked and can remain on the plant for more than a year. The female microsporophyll are more leaf-like than those of other cycads, the greatest similarity to those of the genus Cycas . The scales are flattened at the top, widened and bent up. Each sporophyll carries two, rarely three, ovules. These are straight to the axis (orthotropic), are inclined inwardly to this (inverse) and hang on the axis-facing side of the thickened scales. The opening at the tip of the ovules (micropyle) is very small, leaving only a narrow opening on the Nucellus. The megaspore sheath thickened from 3–4.5 microns in the young ovule to 9-10 microns in the mature seed. The number of archegonia varies between one and ten. The nucleus of the egg is unusually large. The largest female cones of the genus D. spinolosum shows up to 80 centimeters in length and a diameter of up to 30 centimeters.

The seeds are spherical, ovate or ellipsoidal and reach a diameter between about one and four centimeters. They are surrounded by a fleshy, white or cream-colored seed coat. The seed coat consists of three layers, a fleshy outer and inner layer, between which a layer of stone cells lies. The embryo is straight with two cotyledons, these are fused at the top. The embryo carrier (suspensor) is very long and spirally twisted. The seeds are radiated (radiosperm). The germination takes place cryptocotylar, i.e. the cotyledons remain during germination in the seed.

Distribution

Dioon species can be found in Central America between the 15th and 29th northern latitude and are part of the Neotropical floral kingdom. The distribution area includes Honduras and parts of Mexico; it is limited by orographic conditions (height structures) and limited to the north to south mountain ranges in the region. Despite the widespread distribution and ecological plasticity of the genus it is not a vegetation-determining component. The species are found mainly in the tropical rain forest and in tropical wet forests, but also in tropical deciduous rainforests, with softwood rainforests, on dry, rocky slopes, in canyons and on dunes near the coast. [10]

Prehistorical Distribution

At the beginning of the Cenozoic, about 65.5 million years ago, the distribution area of Dioon probably included most of North America. Fossil found in northern Alaska from the Eocene epoch (55.8 - 33.9 Ma) indicate this.

While the climate was still very warm compared to today at the end of the Eocene, around 2.8 million years ago, the early ice age began with the icing of the North Pole, causing the distribution of Dioon to shrink very sharply.

In the Paleocene (66 - 55.8 Ma) Dioon was next to Zamia and Anemia the only today neotropical genus that was still present in the flora of Alaska. During the Tertiary period, about 2.6 million years ago, Dioon died out in almost all of North America with only a few small and isolated relic populations remaining in Central America, where today still the diversity center of this genus lies.

A spread to South America was not possible because the two continents were not connected until about 3 million years ago. After the Isthmus of Panama formed the size of its seeds has been speculated to have been the main obstacle to the further spreading of Dioon into the Caribbean and continental South America.

Ecology

Megasporophyll from Dioon edule with ripe seeds. Right: the dry brown stalk, yellow: two seeds, left: the strongly hairy, flat part of the sporophyll Dioon edule semi02.JPG
Megasporophyll from Dioon edule with ripe seeds. Right: the dry brown stalk, yellow: two seeds, left: the strongly hairy, flat part of the sporophyll

The method of pollination for Dioon plants, whether by wind or animals, has not yet been conclusively determined. The characteristics of the pollen, such as being light, dry, and present in large quantities, suggest wind pollination, which would result in more fertilized ovules near the source of pollen.

However, the arrangement of the ovules in Dioon plants does not support wind pollination, as the pollen has difficulty reaching them and tends to land on the sterile parts of the cone. In contrast, some cycads in the Zamia genus are known to be pollinated by the weevil Rhopalotria mollis . Similarly, in Dioon cones, there is evidence of a large number of beetles from the genus Pharaxonotha , which consume pollen, indicating possible pollination by animals. [11]

Conservation

The genus is particularly at risk from habitat destruction, the International Union for Conservation of Nature (IUCN) lists ten of the eleven species in a vulnerable category on their Red List. One species, D. caputoi, is considered to be critically endangered, three as endangered (D. holmgrenii, D. sonorense and D. tomasellii) and five as vulnerable. (D. califanoi, D. merolae, D. purpusii, D. rzedowskii and D. spinulosum). D. edule is listed as a kind of "near-threatened" list. For D. mejiae, there is not enough data for an assessment of the hazard.

Locations are destroyed primarily by clearing and reclamation of the land. Dioon species are also very popular with international collectors. Plants are dug up and sold. According to a report from the 1992 Washington Convention on Biological Diversity, trade with D. edule individuals reached a level of about 5,800 copies, with D. spinolosum 1,600 - this does not include illicit trade. [12]

Another potential threat to the genus is the lobster louse (Diaspididae) Aulacaspis yasumatsui . The insect originally comes from Southeast Asia and infests there cycads. The pest has now been introduced to Florida, California, India and the UK, where it causes major damage to cycads. There he was also found at D. edule. If the species were to penetrate Central America, it could drastically reduce or even destroy the populations of Dioon in a short time. [13]

Taxonomy

The first description was made in 1843 by John Lindley as Dion. The genus name was given due to the two ovules per megasporophyll: Dioon comes from the ancient Greek prefix δι- di- (= two) and ώόν ōión, ōón (= egg). [14]

The leaf-like character of Sporophylle gives Dioon a special position within the cycads. Dehgan & Dehgan described the genus as the most primitive of the cycads and suggested they put them into their own family. [15] A molecular genetic study of the chloroplast DNA revealed that it is at least the most primitive genus of neotropical cycads. [16]

Classification

Classically, the family of Zamiaceae is divided into two subfamilies with four tribes. Dioon is the only genus in the tribe of the Diooeae, which together with the Encephalarteae forms the subfamily of Encephalartoideae . This classification was based on morphological considerations. A morphological study of relationships in fossil and extant taxa revealed the following cladogram: [17]

A molecular biology analysis of a fragment of 26S ribosomal DNA revealed a somewhat similar cladogram that would dramatically alter the structure of Zamiaceae. [17] Dioon would therefore be the basal genus of the family.

Cycas (Cycadaceae)

Stangeria (Stangeriaceae)

Dioon

Zamia

Encephalartos

Lepidozamia

Ceratozamia

Macrozamia

Microcycas

Species

The genus is commonly divided into two groups by Norstog & Nichols of distinct morphology. The first includes D. mejiae , D. rzedowskii , and D. spinulosum , which are characterized by the greater size of their fronds, trunks, and cones. The second group contains D. califanoi , D. caputoi , D. edule , D. holmgrenii , D. merolae , D. purpusii , D. sonorense , and D. tomasellii , which are smaller overall, with shorter trunks, much shorter fronds, and smaller cones. [18]

A phylogenetic study from 1993 confirmed these clades with one difference - D. caputoi was assigned to the other clade

Phylogeny of Dioon [19] [20]

D. mejiae Standley & Williams

D. rzedowskii De Luca et al.

D. spinulosum Eichl. 1883 ex Dyer

D. edule Lindley

D. merolae De Luca, Sabato & Vázquez-Torres

D. sonorense (De Luca, Sabato & Vázquez-Torres) Chemnick, Gregory & Salas-Morales

D. tomasellii De Luca, Sabato & Vázquez-Torres

D. angustifolium Miquel

D. stevensonii Nicolalde-Morejón & Vovides

D. purpusii Rose

D. argenteum Gregory et al.

D. califanoi De Luca & Sabato

D. caputoi De Luca, Sabato & Vázquez-Torres

D. holmgrenii De Luca, Sabato & Vázquez-Torres

Species accepted: [3]

ImageScientific nameDistribution
Dioon angustifolium furnas 2015.jpg Dioon angustifolium Miq.Mexico (Nuevo León, Tamaulipas)
Dioon argenteum T.J.Greg. & al.Mexico (Oaxaca)
Flickr - brewbooks - Cycads in the greenhouse - Paloma gardens (1).jpg Dioon califanoi De Luca & SabatoMexico (Puebla, Oaxaca)
Cycadales - Dioon caputoi 2.jpg Dioon caputoi De Luca, Sabato & Vázq.TorresMexico (Puebla, Oaxaca)
Dioon edule01.jpg Dioon edule Lindl.Mexico (Nuevo León, Tamaulipas, Veracruz, Guanajuato, Chiapas, Hidalgo, Mexico State, Querétaro, San Luis Potosí)
Dioon holmgrenii 10zz.jpg Dioon holmgrenii De Luca, Sabato & Vázq.TorresMexico (Oaxaca)
Dioon mejae.jpg Dioon mejiae Standl. & L.O.WilliamsHonduras
Dioonmerolae1.jpg Dioon merolae De Luca, Sabato & Vázq.TorresMexico (Oaxaca, Chiapas)
Dioon oaxacensis Gutiérrez-Ortega, Pérez-Farrera & VovidesMexico (Oaxaca)
Dioon planifolium Salas-Morales, Chemnick & GregoryMexico (Oaxaca)
Dioon salas-moralesiae Gutiérrez-Ortega & Pérez-FarreraMexico (Oaxaca)
Dioon purpusii02.jpg Dioon purpusii RoseMexico (Oaxaca)
Dioon rzedowskii.jpg Dioon rzedowskii De Luca, Sabato & Vázq.TorresMexico (Oaxaca)
Dioon sonorense (De Luca, Sabato & Vázq.Torres) Chemnick et al.Mexico (Sonora, Sinaloa)
Dioon spinulosum in Florida.JPG Dioon spinulosum Dyer ex EichlMexico (Veracruz, Oaxaca )
Dioon stevensonii Nic.-Mor. & VovidesMexico (Michoacán, Guerrero)
Dioon tomasellii.jpg Dioon tomasellii De Luca, Sabato & Vázq.TorresMexico (Durango, Jalisco, Nayarit)
Dioon vovidesii Gutiérrez-Ortega & Pérez-FarrerasMexico (Sonora)

Evolution

The earliest fossil finds date back to the Eocene and were found on Kupreanof Island in Alaska. However, it is probable that Dioon dates back to the Jurassic (145.5 - 199.6 ka) or even further, since fossils of other, closely related but more developed cycads were already detected from this era. [21] In the Pleistocene (0.011784-1.8 ka) and Holocene (today -0.011784 ka) eras species have probably carried out several north–south migrations and have followed the warm climatic conditions. [22]

One theory is that the recent dioon species have evolved from a number of vicarious events in the early Cenozoic, with different species representing different ecological conditions. These events must have followed each other for a very short time, which precluded an accumulation of synapomorphies, meaning that not many features could be re-acquired.

The extant species of Dioon are attributed to three prototypes. D. mejiae is considered the most evolutionary advanced species of the genus. D. spinulosum and D. rzedowskii are attributed to a second prototype, which was probably better adapted to mesial conditions, such as areas of medium humidity, which enable them to establish on the Atlantic coast of Oaxaca. The remaining species presumably come from a third, to xerotic, dry areas, adapted to the original form.

Uses

In Honduras, the female cones of D. mejiae are picked to harvest the seeds. These are processed by about 33,000 indigenous people into tamales or tortillas and replace the cornmeal. The fronds are traditionally used on Palm Sunday. [23]

Related Research Articles

<span class="mw-page-title-main">Cycad</span> Division of naked seeded dioecious plants

Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or female. Cycads vary in size from having trunks only a few centimeters to several meters tall. They typically grow slowly and have long lifespans. Because of their superficial resemblance to palms or ferns, they are sometimes mistaken for them, but they are not closely related to either group. Cycads are gymnosperms (naked-seeded), meaning their unfertilized seeds are open to the air to be directly fertilized by pollination, as contrasted with angiosperms, which have enclosed seeds with more complex fertilization arrangements. Cycads have very specialized pollinators, usually a specific species of beetle. Both male and female cycads bear cones (strobili), somewhat similar to conifer cones.

<span class="mw-page-title-main">Gymnosperm</span> Clade of non-flowering, naked-seeded vascular plants

The gymnosperms are a group of seed-producing plants that include conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, and literally means 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, and Ginkgo. The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.

<span class="mw-page-title-main">Conifer cone</span> Reproductive organ on conifers

A conifer cone or pinecone is a seed-bearing organ on gymnosperm plants. It is usually woody, ovoid to globular, including scales and bracts arranged around a central axis, especially in conifers and cycads. The cone of Pinophyta contains the reproductive structures. The woody cone is the female cone, which produces seeds. The male cone, which produces pollen, is usually herbaceous and much less conspicuous even at full maturity. The name "cone" derives from Greek konos, which also gave name to the geometric cone. The individual plates of a cone are known as scales. The umbo of a conifer cone refers, showing up as a protuberance at the end of the two-year-old scale.

<i>Stangeria</i> Monotypic genus of cycads in the family Stangeriaceae

Stangeria eriopus is a cycad endemic to southern Africa. It is the sole species in the genus Stangeria, most closely related to the Australian genus Bowenia, with which it forms the family Stangeriaceae.

<span class="mw-page-title-main">Zamiaceae</span> Family of cycads

The Zamiaceae are a family of cycads that are superficially palm or fern-like. They are divided into two subfamilies with eight genera and about 150 species in the tropical and subtropical regions of Africa, Australia and North and South America.

<i>Zamia</i> Genus of cycads in the family Zamiaceae

Zamia is a genus of cycad of the family Zamiaceae, native to North America from the United States throughout the West Indies, Central America, and South America as far south as Bolivia. 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.

<span class="mw-page-title-main">Sporophyll</span>

A sporophyll is a leaf that bears sporangia. Both microphylls and megaphylls can be sporophylls. In heterosporous plants, sporophylls bear either megasporangia and thus are called megasporophylls, or microsporangia and are called microsporophylls. The overlap of the prefixes and roots makes these terms a particularly confusing subset of botanical nomenclature.

<span class="mw-page-title-main">Bennettitales</span> Extinct order of seed plants

Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales were amongst the most common seed plants of the Mesozoic, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.

<i>Dioon edule</i> Species of cycad

Dioon edule, the chestnut dioon, is a cycad native to Mexico, also known as palma de la virgen. Cycads are among the oldest seed plants and even pre-date the dinosaurs. It belongs to the Zamiaceae plant family within the order Cycadales. The genus name "Dioon" means "two-egged", referring to the two ovules.

<i>Ceratozamia</i> Genus of cycads

Ceratozamia is a genus of New World cycads in the family Zamiaceae. The genus contains 27 known currently living species and one or two fossil species. Most species are endemic to mountainous areas of Mexico, while few species extend into the mountains of Guatemala, Honduras and Belize. The genus name comes from the Greek ceras, meaning horn, which refers to the paired, spreading horny projections on the male and female sporophylls of all species.

A strobilus is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.

<i>Cycas micronesica</i> Species of cycad

Cycas micronesica is a species of cycad found on the island of Yap in Micronesia, the Marianaislands of Guam and Rota, and The Republic of Palau. It is commonly known as federico nut or fadang in Chamorro. The species, previously lumped with Cycas rumphii and Cycas circinalis, was described as a unique species in 1994 by Ken Hill. Paleoecological studies have determined that Cycas micronesica has been present on the island of Guam for about 9,000 years. It has been implicated as a factor in Lytico-Bodig disease, a condition similar to amyotrophic lateral sclerosis (ALS), due to the presence of the neurotoxin BMAA found in its seeds. Seeds were a traditional food source on Guam until the 1960s. The neurotoxin is present due to a symbiosis with cyanobacteria.

<i>Lepidozamia peroffskyana</i> Species of cycad

Lepidozamia peroffskyana is a palm-like cycad in the family Zamiaceae. It is endemic to eastern Australia. The species is named after Count Peroffsky (1794-1857), benefactor of the St. Petersburg Botanical Garden.

<i>Macrozamia glaucophylla</i> Species of plant

Macrozamia glaucophylla is a species of cycad from the genus Macrozamia and the family Zamiaceae. Endemic to New South Wales, Australia, this species has features that resembles palms, although both species are taxonomically quite different. The current population trend of Macrozamia glaucophylla is stable with 2,500 to 10,000 mature individuals. The species are found in several habitats including forest and savanna. Ecologically, Macrozamia glaucophylla lives in terrestrial system, a land-based community of organisms where the biotic and abiotic components interact in the given area.

<i>Macrozamia miquelii</i> Species of cycad

Macrozamia miquelii, is a species of cycad in the plant family Zamiaceae. It is endemic to Queensland and New South Wales in Eastern Australia. Located within sclerophyll forests dominated by eucalyptus trees, the cycad grows on nutrient-poor soils. It is recognised within the Zamiaceae family for its, medium height at 1 m, intermediate size of male and female cones and lighter green leaves compared to other cycads within the plant family of Zamiaceae. The seeds have an orange red sarcotesta which attracts fauna consumption, allowing a mutualistic seed dispersal for the cycad. These seeds are also edible for human consumption if prepared correctly to remove the toxins.

This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary of leaf morphology. For other related terms, see Glossary of phytopathology, Glossary of lichen terms, and List of Latin and Greek words commonly used in systematic names.

<i>Dioon purpusii</i> Species of cycad

Dioon purpusii is a cycad tree endemic to a very small region in Mexico. It is found in Tomellín Canyon and the eastern branches of La Cañada de Cuicatlán.

<i>Dioon tomasellii</i> Species of cycad

Dioon tomasellii is a species of cycad in the family Zamiaceae. It is endemic to Mexico, where it occurs in the states of Durango, Guerrero, Jalisco, Michoacán, and Nayarit.

<i>Zamia integrifolia</i> Species of cycad

Zamia integrifolia, also known as coontie, is a small, tough, woody cycad native to the southeastern United States, the Bahamas, Cuba, the Cayman Islands, and Puerto Rico.

References

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