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''Pandanus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of monocots with about 578 accepted
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. They are palm-like, dioecious trees and shrubs native to the Old World tropics and subtropics. Common names include pandan, screw palm and screw pine. The genus is classified in the order Pandanales, family Pandanaceae, and is the largest in the family.


Description

The species vary in size from small shrubs less than tall, to medium-sized trees tall, typically with a broad canopy, heavy fruit, and moderate growth rate. The trunk is stout, wide-branching, and ringed with many leaf scars. Mature plants can have branches. Depending on the species, the trunk can be smooth, rough, or warty. The roots form a pyramidal tract to hold the trunk. They commonly have many thick stilt roots near the base, which provide support as the tree grows top-heavy with leaves, fruit, and branches. These roots are adventitious and often branched. The top of the plant has one or more crowns of strap-shaped leaves that may be spiny, varying between species from to or longer, and from up to broad. They are dioecious, with male and female flowers produced on different plants. The flowers of the male tree are long and fragrant, surrounded by narrow, white bracts. The female tree produces flowers with round fruits that are also bract-surrounded. The individual fruit is a drupe, and these merge to varying degrees forming multiple fruit, a globule structure, in diameter and have many prism-like sections, resembling the fruit of the pineapple. Typically, the fruit changes from green to bright orange or red as it matures. The fruits can stay on the tree for more than 12 months.


Evolution

The oldest fossil of the genus is ''Pandanus estellae'', which is known from a silicified fruit found in Queensland, Australia, dating to the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
epoch around 32–28 million years ago. Phylogenetic analyses of the maternal DNA of ''Pandanus'' has shown the genus to be divided into two large groups (Clade I and Clade II), with each of those groups further subdivided into two smaller groups (Subclade Ia, Subclade Ib, Subclade IIa, and Subclade IIb). The split between Clades I and II was determined to have occurred during the early Miocene, whereas the splits between Subclade Ia and Ib and between Subclade IIa and IIb were determined to have occurred during the Middle Miocene Climatic Optimum.


Taxonomy

Though often called "pandanus palms", these plants are not closely related to palm trees. The genus is named after the Malay word ''pandan'' given to '' Pandanus amaryllifolius,'' the genus's most commonly known species. The name is derived from Proto-Austronesian *paŋudaN (which became Proto-Malayo-Polynesian *pangdan and Proto-Oceanic *padran). It has many cognates in Austronesian languages, underscoring its importance in Austronesian cultures, including Atayal ''pangran''; Kavalan ''pangzan''; Thao ''panadan''; Tagalog ''pandan''; Chamorro ''pahong''; Manggarai ''pandang''; Malagasy ''fandrana,'' Tongan ''fā''; Tahitian ''fara''; Hawaiian ''hala'' all referring to plants of similar characteristics and/or uses whether in the same genus (particularly '' Pandanus tectorius'') or otherwise (in the case of Māori ''whara'' or ''hara;'' e.g. hara''keke''). ''Pandanus'' is traditionally divided into six subgenera (''Coronata'', ''Kurzia'', ''Lophostigma'', ''Pandanus,'' ''Rykia,'' and ''Vinsonia''), however molecular phylogenetic analyses have shown these subgenera to not represent the true evolutionary history of ''Pandanus'' as all but ''Coronata'' are either polyphyletic or paraphyletic.


Selected species

Note: several species previously placed in ''Pandanus'' subgenera ''Acrostigma'' and ''Martellidendron'' are now in the distinct genera '' Benstonea'' and '' Martellidendron'', respectively. *'' Pandanus aldabraensis'' H.St.John *'' Pandanus amaryllifolius'' Roxb. ex Lindl. *'' Pandanus balfourii'' Martelli *'' Pandanus barkleyi'' Balf.f. *'' Pandanus boninensis'' Warb. *'' Pandanus candelabrum'' P.Beauv. *'' Pandanus carmichaelii'' R.E.Vaughan & Wiehe *'' Pandanus ceylanicus'' Solms *'' Pandanus christmatensis'' Martelli *'' Pandanus clandestinus'' Stone *'' Pandanus conglomeratus'' Balf.f. *'' Pandanus conoideus'' Lam. *'' Pandanus decastigma'' B.C.Stone *'' Pandanus decipiens'' Martelli *'' Pandanus decumbens'' Solms *'' Pandanus drupaceus'' Thouars *'' Pandanus elatus'' Ridl. *'' Pandanus eydouxia'' Balf.f. *'' Pandanus fanningensis'' H.St.John *'' Pandanus forsteri'' C.Moore & F.Muell. *'' Pandanus furcatus'' Roxb. *'' Pandanus gabonensis'' Huynh *'' Pandanus glaucocephalus'' R.E.Vaughan & Wiehe *'' Pandanus grayorum'' *'' Pandanus halleorum'' B.C.Stone *'' Pandanus heterocarpus'' Balf.f. *'' Pandanus iceryi'' Horne ex Balf.f. *'' Pandanus incertus'' R.E.Vaughan & Wiehe *'' Pandanus joskei'' Horne ex Balf.f. *'' Pandanus julianettii'' Martelli *'' Pandanus kaida'' Kurz *'' Pandanus kajui'' Beentje *'' Pandanus lacuum'' H.St.John ex B.C.Stone *'' Pandanus laxespicatus'' Martelli *'' Pandanus livingstonianus'' Rendle *'' Pandanus leram'' *'' Pandanus microcarpus'' Balf.f. *'' Pandanus montanus'' Bory *'' Pandanus multispicatus'' Balf.f. *'' Pandanus odorifer'' ( Forssk.) Kuntze *'' Pandanus obeliscus'' *'' Pandanus palustris'' Thouars *'' Pandanus parvicentralis'' Huynh *'' Pandanus prostratus'' Balf.f. *'' Pandanus pyramidalis'' Barkly ex Balf.f. *'' Pandanus rigidifolius'' R.E.Vaughan & Wiehe *'' Pandanus sechellarum'' Balf.f. *'' Pandanus spathulatus'' Martelli *'' Pandanus spiralis'' R.Br. *'' Pandanus tectorius'' Parkinson ex Du Roi *'' Pandanus tenuifolius'' Balf.f. *'' Pandanus teuszii'' Warb. *'' Pandanus thomensis'' Henriq. *'' Pandanus tonkinensis'' B.C.Stone *'' Pandanus utilis'' Bory *'' Pandanus vandermeeschii'' Balf.f. *'' Pandanus verecundus'' Stone


Distribution and habitat

''Pandanus'' is a Paleotropical genus. The greatest number of species are found in
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
and Malaysia.


Ecology

These plants grow from sea level to an altitude of . Pandanus trees are of cultural, health, and economic importance in the Pacific, second only to the coconut on atolls. They grow wild mainly in semi-natural vegetation in littoral habitats throughout the tropical and subtropical Pacific, where they can withstand drought, strong winds, and salt spray. They propagate readily from seed, but popular cultivars are also widely propagated from branch cuttings by local people. Species growing on exposed coastal headlands and along beaches have thick 'stilt roots' as anchors in the loose sand. Those stilt roots emerge from the stem, usually close to but above the ground, which helps to keep the plants upright and secure them to the ground. While ''Pandanus'' are distributed throughout the tropical and subtropical islands and coastlines of the Atlantic, Indian and Pacific Oceans, they are most numerous on the low islands and barren atolls of Polynesia and Micronesia. Other species are adapted to mountain habitats and riverine forests. The tree is grown and propagated from shoots that form spontaneously in the axils of lower leaves. Pandanus fruits are eaten by animals including bats, rats, crabs, and elephants, but the vast majority of species are dispersed primarily by water. Its fruit can float and spread to other islands without help from humans. Climate and soil analyses based on a molecular phylogenetic framework have shown that the largest split in ''Pandanus'' (between Clades I and II) is also associated with a difference in climate and soil conditions between the two groups. Clade II ''Pandanus'' species are associated with more seasonal
temperature Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...
and precipitation regimes, and more well-draining soils, than Clade I ''Pandanus'' species. In addition, this split is also associated with the occurrence of specialized layer of water storage tissue (also known as hydrenchyma) in Clade II species of ''Pandanus'' that is thought to aid in their adaptation to more water-stressed environments.


Uses

''Pandanus'' has multiple uses, which is dependent in part on each type and location. Some ''Pandanus'' are a source of food, while others provide raw material for clothing, basket weaving and shelter. ''Pandanus'' leaves are used for handicrafts. Artisans collect the leaves from plants in the wild, cutting only mature leaves so that the plant will naturally regenerate. The leaves are sliced into fine strips and sorted for further processing. Weavers produce basic pandan mats of standard size or roll the leaves into pandan ropes for other designs. This is followed by a coloring process, in which pandan mats are placed in drums with water-based colors. After drying, the colored mats are shaped into final products, such as placemats or jewelry boxes. Final color touch-ups may be applied. The species in Hawaiʻi are called ''hala'', and only the dry leaves ''(lauhala)'' are collected and used for Lauhala weaving. Traditions of weaving pandanus to source fabric material were widespread among Polynesians even as they migrated reaching colder latitudes (like the islands of New Zealand) where no pandanus grew, which later Māori generations simply adapted their skills with native plants like '' Phormium'' having superficially similar properties, even reflected in their names (e.g. the aforementioned '' harakeke'', and '' wharariki''). ''Pandanus'' leaves from '' Pandanus amaryllifolius'' are used widely in Southeast Asian and South Asian cuisines to add a distinct aroma to various dishes and to complement flavors like chocolate. Because of their similarity in usage, pandan leaves are sometimes referred to as the "vanilla of Asia." Fresh leaves are typically torn into strips, tied in a knot to facilitate removal, placed in the cooking liquid, then removed at the end of cooking. Dried leaves and bottled extract may be bought in some places. Finely sliced pandan leaves are used as fragrant confetti for Malay weddings, graves etc. Pandan leaves are known as ''Daun pandan'' in Indonesian and Malaysian Malay; ''Dahon ng pandan'' () or simply ''pandan'' in Filipino; 斑蘭 (''bān lán'') in Mandarin; as ใบเตย (''bai toei''; ) in Thai, ''lá dứa'' in Vietnamese; ''pulao data'' in Bengali; and ''rampe'' in Sinhalese and Hindi. In India, particularly in Nicobar Islands, pandanus fruit is staple food of Shompen people and Nicobarese people. In Sri Lanka, pandan leaves are used heavily in both vegetable and meat dishes and are often grown in homes. It is common practice to add a few pieces of pandan leaf when cooking red or white rice as well. In Southeast Asia, pandan leaves are mainly used in sweets such as coconut jam and pandan cake. In Indonesia and Malaysia, pandan is also added to rice and curry dishes such as '' nasi lemak''. In the Philippines, pandan leaves are commonly paired with coconut meat (a combination referred to as ''buko pandan'') in various desserts and drinks like '' maja blanca'' and '' gulaman''. In Indian cooking, the leaf is added whole to biryani, a kind of rice pilaf, made with ordinary rice (as opposed to that made with the premium-grade basmati rice). The basis for this use is that both basmati and pandan leaf contains the same aromatic flavoring ingredient, 2-acetyl-1-pyrroline. In
Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...
, pandan leaves are a major ingredient used in the country's cuisine. '' Kewra'' (also spelled ''Kevda'' or ''Kevada'') is an extract distilled from the pandan flower, used to flavor drinks and desserts in Indian cuisine. Also, ''kewra'' or ''kevada'' is used in religious worship, and the leaves are used to make hair ornaments worn for their fragrance as well as decorative purpose in western India. Species with large and medium fruit are edible, notably the many cultivated forms of '' P. tectorius'' (''P. pulposus'') and '' P. utilis''. The ripe fruit can be eaten raw or cooked, while partly ripe fruit should be cooked first. Small-fruited pandanus may be bitter and astringent. Karuka nuts (''P. julianettii'') are an important staple food in New Guinea. Over 45 cultivated varieties are known. Entire households will move, and in some areas will speak a pandanus language at harvest time. The taste is like coconut or walnuts. Throughout Oceania, almost every part of the plant is used, with various species different from those used in Southeast Asian cooking. ''Pandanus'' trees provide materials for housing; clothing and textiles including the manufacture of dilly bags (carrying bags), fine mats or '' ie toga''; sails, food, medication, decorations, fishing, and religious uses. In the Vanuatu Archipelago, natives make woven fish traps from the hard interior root of the pandanus, made like a cage having a narrow entrance.SLICE documentary, , The Isle of Futuna / April 2022, minutes 18:39–ff.


See also

* * – vessels of the Caroline Islands which traditionally had pandanus mat sails *


References


Further reading


Germplasm Resources Information Network: ''Pandanus''
*Wagner, W. L., Herbst, D. R., & Sohmer, S. H. (1990). ''Manual of the flowering plants of Hawaiʻi''. * photos and text by Dave Kimble * - photo essay

* ttps://www.youtube.com/watch?v=0Df1HYtTPv4 Pandanus simplex fruit eaten by Varanus olivaceus, Polillo Island, Philippines. br>"Hala: The Hawaiian Aphrodisiac"
Article by Shannon Wianecki describing Hawaiian cultural uses for pandanus. '' Maui No Ka 'Oi Magazine'' Volume 15 Number. 1 (Jan 2011).


External links

*
Biological Analytics of Pandan
{{Authority control Pandanales genera Medicinal plants