You can help expand this article with text translated from the corresponding article in Spanish. (January 2016)Click [show] for important translation instructions.
|
Pandanales Temporal range: Mid Cretaceous – Recent | |
---|---|
Carludovica palmata | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Monocots |
Order: | Pandanales R.Br. ex Bercht. & J.Presl [1] |
Type species | |
Pandanus tectorius Parkinson | |
Families | |
Pandanales, the pandans or screw-pines, is an order of flowering plants placed in the monocot clade in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web systems. Within the monocots Pandanales are grouped in the lilioid monocots where they are in a sister group relationship with the Dioscoreales. Historically the order has consisted of a number of different families in different systems but modern classification of the order is based primarily on molecular phylogenetics despite diverse morphology which previously placed many of the families in other groupings based on apparent similarity. Members of the order have a subtropical distribution and includes trees, shrubs, and vines as well as herbaceous plants. The order consists of 5 families, 36 genera and about 1,610 species.
Pandanales are highly diverse including large arboraceous plants of tropical rainforests and coastal areas, climbing vines and lianas, as well as very small achlorophyllous (mycoheterotrophic) and saprophytic herbaceous forest floor species. This has made it difficult to reliably define synapomorphies, but the loss of trimery distinguishes many of them from other lilioid monocots. [2]
The Pandanales order is distinctive with its highly variable and hardly definable floral morphology, [2] especially the number of stamens and their structure as well as many other characteristics. In some of the members, different interpretations exist regarding the composition and organization of the reproductive structures.
The order includes plants with traits that seem atypical when compared to other groups of monocots. A good example is the female reproductive organ and its position relative to other parts of the flower. Some of the species included in the families Pandanaceae and Stemonaceae show flowers formed from only one carpel, while in the Triuridaceae, a family that lacks chlorophyll, the carpels are free from each other. In fact, the Triuridaceae possess the least "typical" flower morphology in the order.[ citation needed ]
The components of the order sensu APG have been difficult to place consistently, and historically have been associated with a number of other groupings. [2] The Bentham & Hooker system (1883) had a similar order under the name Nudifloreae, incorporating:
The Wettstein system (1935) placed the order in class Monocotyledones and used a different circumscription, incorporating:
The Cronquist system (1981) placed the order in subclass Arecidae in class Liliopsida [=monocotyledons] with only one family:
In the classification system of Dahlgren the Pandanales were in the superorder Pandaniflorae (also called Pandananae) with the single family Pandanaceae.
The APG III system (2009) places the Pandanales in the monocots. Both the APG III and APG II systems include five families in this order. [1] Since the morphology of the order varies on such scale, its classification and phylogeny are based on genetic analyses.
Inside the order, some doubt remains about the position of the entirely mycoheterotrophic family Triuridaceae, since it is the only one on which genetic analyses have not yet been applied. With high probability, the family may be sister to the Velloziaceae, but similarities with the family Zingiberaceae (which is a part of a whole different order – Zingiberales) do not exclude the chance for a different phylogeny. The family Velloziaceae on its own is placed at the base of the tree. The Pandanaceae and Cyclanthaceae are sister groups, and they form a clade which on its own is sister to the Stemonaceae (a family composed of two more clades).
The order Dioscoreales holds sister relationship with Pandanales by diverging from them around 121 million years ago in the mid-Cretaceous. The formation of the crown groups took place with a difference of 2 million years between the orders - 116 Mya for the Dioscoreales and 114 Mya for the Pandanales. However, the stem group of the Pandanales is much older and goes back to 130 Mya in the early Cretaceous. [3]
| ||||||||||||||||||||||||||||
The composition of the order in APG III and APG II were slightly from that in the 1998 APG system, which used the circumscription [1]
APG III includes the following families;
The order contains members mainly distributed in all the tropical and subtropical regions of the world, both New and Old World (including Africa, South America, Australia and Asia). [2] Also, a species is present in China.
The species are members of various ecological groups, including tropical shrubs, lianas and trees, xerophytic plants, mycoheterotrophs, as well as different herbaceous representatives.
Several species in this order produce strap-like leaves used for basketry and mats; Pandanus (Pandanaceae) is used across Oceania for thatch, basketry, and to make cloth, and Carludovica palmata (Cyclanthaceae) leaves are made into Panama hats. Other members as Stemona are present in traditional Chinese medicine and possess medical properties. Some species are used as insecticides. Some species of Pandanus are used as ornamentals. Pandanus amaryllifolius is a common food flavoring in Southeast Asia and karuka (Pandanus julianettii) is a staple food in New Guinea.
Asparagales is an order of plants in modern classification systems such as the Angiosperm Phylogeny Group (APG) and the Angiosperm Phylogeny Web. The order takes its name from the type family Asparagaceae and is placed in the monocots amongst the lilioid monocots. The order has only recently been recognized in classification systems. It was first put forward by Huber in 1977 and later taken up in the Dahlgren system of 1985 and then the APG in 1998, 2003 and 2009. Before this, many of its families were assigned to the old order Liliales, a very large order containing almost all monocots with colorful tepals and lacking starch in their endosperm. DNA sequence analysis indicated that many of the taxa previously included in Liliales should actually be redistributed over three orders, Liliales, Asparagales, and Dioscoreales. The boundaries of the Asparagales and of its families have undergone a series of changes in recent years; future research may lead to further changes and ultimately greater stability. In the APG circumscription, Asparagales is the largest order of monocots with 14 families, 1,122 genera, and about 36,000 species.
The Dioscoreales are an order of monocotyledonous flowering plants, organized under modern classification systems, such as the Angiosperm Phylogeny Group or the Angiosperm Phylogeny Web. Among monocot plants, Dioscoreales are grouped with the lilioid monocots, wherein they are a sister group to the Pandanales. In total, the order Dioscoreales comprises three families, 22 genera and about 850 species.
Liliales is an order of monocotyledonous flowering plants in the Angiosperm Phylogeny Group and Angiosperm Phylogeny Web system, within the lilioid monocots. This order of necessity includes the family Liliaceae. The APG III system (2009) places this order in the monocot clade. In APG III, the family Luzuriagaceae is combined with the family Alstroemeriaceae and the family Petermanniaceae is recognized. Both the order Lililiales and the family Liliaceae have had a widely disputed history, with the circumscription varying greatly from one taxonomist to another. Previous members of this order, which at one stage included most monocots with conspicuous tepals and lacking starch in the endosperm are now distributed over three orders, Liliales, Dioscoreales and Asparagales, using predominantly molecular phylogenetics. The newly delimited Liliales is monophyletic, with ten families. Well known plants from the order include Lilium (lily), tulip, the North American wildflower Trillium, and greenbrier.
Monocotyledons, commonly referred to as monocots, are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of the major groups into which the flowering plants have traditionally been divided; the rest of the flowering plants have two cotyledons and are classified as dicotyledons, or dicots.
Dioscoreaceae is a family of monocotyledonous flowering plants, with about 715 known species in nine genera. The best-known member of the family is the yam.
Petrosaviaceae is a family of flowering plants belonging to a monotypic order, Petrosaviales. Petrosaviales are monocots, and are grouped within the lilioid monocots. Petrosaviales is a very small order composed of one family, two genera and four species accepted in 2016. Some species are photosynthetic (Japonolirion) and others are rare, leafless, chlorophyllous, mycoheterotrophic plants (Petrosavia). The family is found in low-light montane rainforests in Japan, China, Southeast Asia and Borneo. They are characterised by having bracteate racemes, pedicellate flowers, six persistent tepals, septal nectaries, three almost-distinct carpels, simultaneous microsporogenesis, monosulcate pollen, and follicular fruit.
Burmanniaceae is a family of flowering plants, consisting of 99 species of herbaceous plants in eight genera.
Asphodelaceae is a family of flowering plants in the order Asparagales. Such a family has been recognized by most taxonomists, but the circumscription has varied widely. In its current circumscription in the APG IV system, it includes about 40 genera and 900 known species. The type genus is Asphodelus.
Pandanaceae is a family of flowering plants native to the tropics and subtropics of the Old World, from West Africa through the Pacific. It contains 982 known species in five genera, of which the type genus, Pandanus, is the most important, with species like Pandanus amaryllifolius and karuka being important sources of food. The family likely originated during the Late Cretaceous.
The APG system of plant classification is the first version of a modern, mostly molecular-based, system of plant taxonomy. Published in 1998 by the Angiosperm Phylogeny Group, it was replaced by the improved APG II in 2003, APG III system in 2009 and APG IV system in 2016.
Triuridaceae are a family of tropical and subtropical flowering plants, including nine genera with a total of approximately 55 known species. All members lack chlorophyll and are mycoheterotrophic. The heterotrophic lifestyle of these plants has resulted in a loss of xylem vessels and stomata, and a reduction of leaves to scales.
Cyclanthaceae is a family of flowering plants.
Velloziaceae is a family of monocotyledonous flowering plants. The APG II system, of 2003, also recognizes this family, and assigns it to the order Pandanales.
The Stemonaceae are a family of monocotyledonous flowering plants placed in the order Pandanales. The family consists of four genera with ca 37 known species distributed in areas with seasonal climate across Southeast Asia and tropical Australia. One native species is found in the United States. In earlier systems the family was called Roxburghiaceae, after Roxburghia, now Stemona.
The Kubitzki system is a system of plant taxonomy devised by Klaus Kubitzki, and is the product of an ongoing survey of vascular plants, entitled The Families and Genera of Vascular Plants, and extending to 15 volumes in 2018. The survey, in the form of an encyclopedia, is important as a comprehensive, multivolume treatment of the vascular plants, with keys to and descriptions of all families and genera, mostly by specialists in those groups. The Kubitzki system served as the basis for classification in Mabberley's Plant-Book, a dictionary of the vascular plants. Mabberley states, in his Introduction on page xi of the 2008 edition, that the Kubitzki system "has remained the standard to which other literature is compared".
Arecidae is a botanical name at the rank of subclass. Circumscription of the subclass will vary with the taxonomic system being used ; the only requirement being that it includes the family Arecaceae.
Thismiaceae is a family of flowering plants whose status is currently uncertain. The Angiosperm Phylogeny Group classifications merge Thismiaceae into Burmanniaceae, noting that some studies have suggested that Thismiaceae, Burmanniaceae and Taccaceae should be separate families, whereas others support their merger.
The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a further revision, the APG IV system.
Lilioid monocots is an informal name used for a grade of five monocot orders in which the majority of species have flowers with relatively large, coloured tepals. This characteristic is similar to that found in lilies ("lily-like"). Petaloid monocots refers to the flowers having tepals which all resemble petals (petaloid). The taxonomic terms Lilianae or Liliiflorae have also been applied to this assemblage at various times. From the early nineteenth century many of the species in this group of plants were put into a very broadly defined family, Liliaceae sensu lato or s.l.. These classification systems are still found in many books and other sources. Within the monocots the Liliaceae s.l. were distinguished from the Glumaceae.