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Pandanus
Pandanus
Pandanus
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Not to be confused with Pandalus.

Pandanus
Temporal range: Oligocene-Recent 30–0 Ma
Fruit of Pandanus utilis
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Pandanales
Family: Pandanaceae
Genus: Pandanus
Parkinson[1]
Species

See List of Pandanus species

Synonyms[1]
20 synonyms
  • Athrodactylis J.R.Forst. & G.Forst.
  • Barrotia Gaudich.
  • Bryantia Webb ex Gaudich.
  • Doornia de Vriese
  • Dorystigma Gaudich.
  • Eydouxia Gaudich.
  • Foullioya Gaudich.
  • Hasskarlia Walp.
  • Heterostigma Gaudich.
  • Hombronia Gaudich.
  • Jeanneretia Gaudich.
  • Keura Forssk.
  • Marquartia Hassk.
  • Pandanus Rumph. ex L.f.
  • Roussinia Gaudich.
  • Rykia de Vriese
  • Souleyetia Gaudich.
  • Sussea Gaudich.
  • Tuckeya Gaudich.
  • Vinsonia Gaudich.

Pandanus is a genus of monocots with about 578 accepted species.[1] 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,[2][3] and is the largest in the family.[4]

Description

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Aerial, prop roots[5]

The species vary in size from small shrubs less than 1 metre (3+1⁄2 feet) tall, to medium-sized trees 20 m (66 ft) tall, typically with a broad canopy, heavy fruit, and moderate growth rate.[6][self-published source?] The trunk is stout, wide-branching, and ringed with many leaf scars.[7] Mature plants can have branches.[7] Depending on the species, the trunk can be smooth, rough, or warty.[8] The roots form a pyramidal tract to hold the trunk.[9] They commonly have many thick stilt roots near the base, which provide support as the tree grows top-heavy with leaves, fruit, and branches.[10] 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,[2][3] varying between species from 30 centimetres (12 inches) to 2 m (6+1⁄2 ft) or longer, and from 1.5 cm (5⁄8 in) up to 10 cm (4 in) broad.

They are dioecious, with male and female flowers produced on different plants. The flowers of the male tree are 2–3 cm (3⁄4–1+1⁄4 in) 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, 10–20 cm (4–8 in) 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

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The oldest credible fossil of the genus is Pandanus estellae, which is known from a silicified fruit found in Queensland, Australia, dating to the Oligocene epoch around 32–28 million years ago.[11] 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).[12][13] 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.[13]

Taxonomy

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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.[14] 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. harakeke).[15][16] 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.[12][13]

Selected species

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Main article: List of Pandanus species
Pandanus repens
Pandanus simplex

Note: several species previously placed in Pandanus subgenera Acrostigma and Martellidendron are now in the distinct genera Benstonea and Martellidendron, respectively.[12]

Distribution and habitat

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Pandanus is a Paleotropical genus.[4] The greatest number of species are found in Madagascar and Malaysia.[17][page needed]

Ecology

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These plants grow from sea level to an altitude of 3,300 m (10,800 ft). Pandanus trees are of cultural, health, and economic importance in the Pacific, second only to the coconut on atolls.[18][19] 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.[2]

Species growing on exposed coastal headlands and along beaches have thick 'stilt roots' as anchors in the loose sand.[2][20] 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.[10]

While Pandanus are distributed throughout the tropical and subtropical islands and coastlines of the Atlantic, Indian and Pacific Oceans,[21][22][23] they are most numerous on the low islands and barren atolls of Polynesia and Micronesia.[24][25][26][27] Other species are adapted to mountain habitats and riverine forests.[28]

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.[9] Its fruit can float and spread to other islands without help from humans.[22]

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.[13] Clade II Pandanus species are associated with more seasonal temperature and precipitation regimes, and more well-draining soils, than Clade I Pandanus species.[13] 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.[13]

Uses

[edit]

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).[29]

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."[30][31][32] 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.[citation needed]

Pandan leaves are known as Daun pandan in Indonesian and Malaysian Malay; Dahon ng pandan (lit. 'pandan leaf') or simply pandan in Filipino; 斑蘭 (bān lán) in Mandarin; as ใบเตย (bai toei; pronounced [bāj.tɤ̄ːj]) 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.[33]

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.[34]

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, pandan leaves are a major ingredient used in the country's cuisine.[35]

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.[24]

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,[36] while partly ripe fruit should be cooked first.[37] Small-fruited pandanus may be bitter and astringent.[36]

Karuka nuts (P. julianettii) are an important staple food in New Guinea.[38] Over 45 cultivated varieties are known.[39] Entire households will move,[40] and in some areas will speak a pandanus language at harvest time.[41][42] The taste is like coconut[38][39][43] or walnuts.[44]

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 ʻie toga; sails,[45] food, medication,[citation needed] 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.[46]

See also

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References

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Further reading

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Pandanus

View on Grokipedia
from Grokipedia
Pandanus is a of monocotyledonous flowering in the family , consisting of over 500 accepted of palm-like, dioecious trees, shrubs, and climbers characterized by their stout trunks supported by distinctive stilt roots, long strap-shaped leaves arranged in a spiral pattern, and compound inflorescences producing drupaceous fruits. These typically range from 1 to 20 meters in height and are adapted to a variety of tropical environments, with leaves often featuring sharp, toothed margins that give the genus its common name, "screw pine." Taxonomically, Pandanus belongs to the order and is the largest genus in the family, which includes five genera in total. The genus is distributed across the tropics and subtropics, from tropical and through , , and the islands of the Indian and Pacific Oceans, including and , where species diversity is highest in regions like and . They inhabit diverse ecosystems such as coastal areas, riverbanks, forests, and even altitudes up to 3,300 meters, often thriving in unstable, muddy, or anaerobic soils due to their prop roots that aid in . While some species extend to , the genus is absent from the . Morphologically, Pandanus species exhibit significant variation, with leaves reaching lengths of 30 cm to 2 meters, arranged in three ranks to form a screw-like rosette, and trunks often bearing sharp prickles. Reproduction is primarily sexual, with flowers on separate , though some species reproduce asexually via ; seeds are dispersed by animals in forested habitats or by along coastlines. Ecologically, these play key roles in , providing windbreaks and natural fences in coastal and riparian zones. In human cultures, particularly in the Pacific Islands, Asia, and Africa, Pandanus holds significant economic and traditional value, with leaves widely used for weaving mats, hats, baskets, thatch, and cordage due to their fibrous strength. Certain species, such as Pandanus tectorius, produce edible fruits that are nutritious and form part of local diets, while others like Pandanus amaryllifolius are cultivated for their fragrant leaves used in flavoring food, such as rice and desserts. Additionally, some species yield oils from seeds and contribute to medicinal practices, underscoring their cultural importance in indigenous communities.

Description and Morphology

Physical Characteristics

Pandanus species exhibit a distinctive palm-like growth habit, forming stout, unbranched or sparsely branched trees or shrubs that can reach heights of 1 to 20 meters, with canopy spreads varying from 4 to 14 meters or more depending on the . These plants are dioecious, with separate male and female individuals, and belong to the family. The stems are typically grayish or reddish-brown, smooth to flaky, and marked by prominent leaf scars and occasional prickles, providing structural support in tropical environments. Morphological characteristics vary widely among the approximately 750 , with some reaching 20 m and others remaining shrub-like under 1 m. A key morphological feature is the extensive system of prop roots or stilt roots, which emerge from the lower portion of the stem, usually close to but above the ground, and extend into the , anchoring the plant in unstable substrates like sandy beaches or wetlands. Leaves are spirally arranged in three vertical rows at the branch apices, linear to sword-shaped, reaching up to 3 meters in length and 3 to 16 centimeters wide in larger species, with parallel venation, a conspicuous midrib, and often spiny margins and midribs featuring prickles. These adaptations enhance stability and deter herbivores in coastal and forested habitats. Male inflorescences consist of catkin-like or racemes with tiny, fragrant white or yellow flowers, often subtended by large white bracts, while female inflorescences form dense globose heads that develop into syncarps. The fruits are aggregate structures resembling pineapples, varying from 10 to 30 centimeters long and up to 20 centimeters in diameter across , composed of numerous wedge-shaped drupelets forming ovoid to globose heads, with colorful or orange exteriors and buoyant properties that facilitate water dispersal. The mesocarp is fibrous and fleshy, enclosing hard, bony endocarps with obovoid seeds.

Reproduction

Pandanus species are dioecious, with male and female reproductive structures on separate plants, enabling specialized . Male inflorescences produce copious amounts of lightweight , which is primarily dispersed by wind in many species such as , facilitating of female flowers. However, recent research on Pandanus odorifer has identified sap beetles (Amystrops spp.) as effective pollinators, suggesting may play a role in at least some fragrant screw pine species, overturning prior assumptions of exclusive anemophily. Following successful , female flowers develop into syncarpous fruits—fused aggregates of multiple drupes—each containing numerous seeds embedded within a fibrous, buoyant structure. Fruit dispersal in Pandanus occurs mainly through hydrochory, as the lightweight, air-filled syncarps float on surfaces and are carried by currents or rivers, aiding long-distance in coastal and environments. Some species exhibit zoochory, where animals consume or transport the fruits, combining flotation with biotic assistance for broader dissemination. germination typically requires moist, well-drained conditions; the seeds, encased in hard endocarps, emerge via a basal germination tube after or prolonged soaking to break , often taking 1–1.5 months. Asexual reproduction in Pandanus occurs vegetatively through basal suckers or stem cuttings, particularly in cultivated species such as Pandanus amaryllifolius, which is sterile and relies entirely on these methods for propagation. Suckers emerge from the rootstock and can be separated to establish new plants, while cuttings of lateral shoots root readily in humid conditions. Flowering phenology is seasonal in many tropical Pandanus species, with female plants blooming 1–3 times annually and males more frequently, often triggered by rainfall patterns or photoperiod changes in humid lowlands.

Taxonomy and Classification

History of Classification

The genus Pandanus was established by in his in 1753, where he described the Pandanus odoratissimus L. based on earlier accounts from Southeast Asian flora. The name derives from the Malay word "pandan," referring to the fragrant qualities of certain species, particularly Roxb., which became widely recognized for its aromatic leaves used in culinary and cultural contexts. In the early , Robert Brown formalized the family in 1810 within his Prodromus Florae Novae Hollandiae, classifying Pandanus alongside related genera based on shared morphological traits such as prop roots and aggregated fruits, marking a shift from Linnaean groupings toward a more natural system. Brown's work highlighted the distribution and dioecious nature of the genus, drawing from collections during his Australian expeditions. Throughout the , classifications refined Pandanus within , emphasizing infrageneric divisions based on fruit structure and leaf anatomy, though comprehensive subgeneric schemes emerged later. In the , Harold St. John undertook extensive revisions through his multi-part series Revision of the Genus Pandanus Stickman (1960–1988), proposing over 1,000 species across numerous sections and subgenera, often based on colonial-era specimens from the Pacific and . These proliferations were later contested for over-splitting due to insufficient differentiation, leading to widespread synonymy as subsequent botanists consolidated taxa using more rigorous morphological and distributional criteria. Modern taxonomy accepts approximately 600–750 species in Pandanus, with 563 currently accepted per Plants of the World Online (as of 2025), reflecting ongoing revisions following the 2012 circumscription by Callmander et al., who estimated around 450 species after elevating subgenus Acrostigma to the separate genus Benstonea based on integrated morphological and preliminary molecular data. Recent phylogenetic studies, such as the 2025 analysis of complete plastomes from 50 Pandanus species, have used high-resolution DNA markers to reassess subgeneric boundaries, confirming non-monophyly in traditional groupings like subgenus Pandanus and Acanthostigma while linking morphological adaptations (e.g., water-storage tissues) to climatic niches in the Paleotropics; the study indicates that subgeneric classification requires revision. These efforts employ chloroplast genomes to resolve polytomies unresolved by earlier nuclear and plastid markers, providing a framework for ongoing revisions. Classification challenges persist due to extreme morphological variation within species complexes, particularly in leaf spininess and fruit aggregation, which has fueled historical synonymy rates exceeding 50% in some regions. Incomplete type specimens from 19th- and early 20th-century colonial collections, often lacking locality details or reproductive structures, further complicate verification and phylogenetic placement.

Species Diversity

The genus Pandanus comprises approximately 600–750 species according to recent assessments, with 563 accepted per (as of 2025), though estimates account for ongoing taxonomic revisions and undescribed . The genus is divided into eight subgenera, including Pandanus, Rykia, Lophostigma, Kurzia, Coronata, Vinsonia, and others, as established in foundational infrageneric classifications, though recent studies suggest revisions are needed. High species diversity occurs in , particularly in (encompassing , the , and ), where the greatest concentration of species is documented, alongside notable in the Pacific islands featuring insular specialists such as Pandanus tectorius. Notable examples include Pandanus amaryllifolius (native to ), Pandanus utilis (widespread in the ), and Pandanus spiralis (endemic to ). Infrageneric variation within Pandanus is pronounced, with species identification relying on morphological keys such as the arrangement and density of leaf spines (e.g., marginal versus intramarginal types) and fruit morphology, including syncarp size, drupelet fusion, and coloration patterns. Recent taxonomic adjustments include the 2012 segregation of the genus Benstonea from Pandanus sensu lato, transferring approximately 50 species based on distinct morphological traits like peduncle structure and molecular phylogenetic evidence, thereby refining the circumscription of the core Pandanus genus.

Evolutionary History

Origins and Phylogeny

The family is estimated to have originated in the , approximately 79–133 million years ago, with analyses suggesting a crown age of 26–35 million years ago based on bi-organellar phylogenomic data. Although early hypotheses proposed a Gondwanan origin supported by distributions across southern continents, recent biogeographic reconstructions favor a Laurasian cradle followed by dispersals to former Gondwanan landmasses like , , and during the Late Eocene. The genus Pandanus within diverged in the Paleotropics around 7–21 million years ago, during the , as inferred from plastid and mitochondrial divergence estimates. Phylogenetic analyses using molecular markers such as rbcL and matK genes confirm Pandanus as monophyletic, positioned as sister to the Freycinetia-Martellidendron pair within , with Sararanga basal to the family. Long-distance dispersal, rather than vicariance, is the dominant mechanism for the Pacific colonization of Pandanus, as evidenced by the P. tectorius complex originating in eastern within the last 6 million years and radiating via ocean currents. Key evolutionary adaptations include the development of prop roots for anchorage in unstable substrates and buoyant, multi-seeded fruits facilitating hydrochorous dispersal and island-hopping across paleotropical archipelagos. Subgeneric phylogeny reveals basal clades centered in Africa and Asia, with derived insular radiations in the Indo-Pacific; plastome sequencing of 50 species identifies two major clades and four subclades, with only subgenus Coronata monophyletic, necessitating taxonomic revisions. Recent 2025 phylogenomic studies correlate morphological traits, such as hypertrophied water-storage tissue in clade II, with climatic niches involving seasonal precipitation and well-draining soils, linking early Miocene clade divergences to the onset of the Southeast Asian monsoon. Evidence of hybridization is rare but documented in sympatric populations, such as in Queensland where overlapping species ranges suggest gene flow influencing species boundaries, as supported by observations of potential interspecific crosses.

Fossil Record

The fossil record of Pandanus is sparse, primarily due to the poor preservation potential of monocotyledonous plants, with most evidence consisting of grains, leaf impressions, and rare fruits. The earliest macrofossils attributable to the date to the Eocene epoch, around 50 million years ago, including leaf impressions identified as Pandanus eocenicus from early Eocene sediments in , . Pollen records of extend slightly further back, with comparable forms from Palaeocene deposits in and , suggesting an early presence on Laurasian and former Gondwanan landmasses. No fossils predate the , when the oldest Pandanaceae pollen appears in Maastrichtian sediments from . Oligocene and Miocene records provide additional insights into the diversification of prop root-bearing structures characteristic of modern Pandanus. A notable example is the silicified fruit of the extinct species Pandanus estellae from deposits in central Queensland, Australia, representing the oldest confirmed of the genus and indicating early adaptation to coastal environments. In Southeast Asia, Miocene pollen attributed to Pandanus has been documented from various sites, including syncarp-like impressions that hint at the development of aggregated structures, though direct amber-preserved examples remain elusive. Fossil distributions include occurrences in former Gondwanan landmasses such as , , and , consistent with early long-distance dispersals from a Laurasian origin following the family's diversification. These fossils align with biogeographic models emphasizing long-distance dispersal to account for the Paleotropical expansion of Pandanus lineages, rather than vicariance associated with the breakup. The limited record, however, reflects challenges in fossilizing delicate monocot tissues like leaves and roots, potentially underestimating the true antiquity and diversity. Recent discoveries in the 2020s, including pollen analyses from sediments in the Pacific (e.g., New Ireland, ), document expansions of Pandanus populations, likely influenced by sea-level changes and human-mediated dispersal.

Distribution and Habitat

Geographic Range

The genus Pandanus is native to the tropical and subtropical regions of the , with its westernmost limit in , where species such as Pandanus candelabrum occur from to . The distribution extends eastward across , the islands, and , southern , , , and , reaching as far as the Pacific islands including , , , Pitcairn, and Henderson Islands. This broad Paleotropical range spans from coastal lowlands to inland areas, encompassing approximately 600–700 species adapted to diverse island and continental environments. Centers of diversity for Pandanus are concentrated in the Indo-Malayan region, particularly and , and in , where a significant proportion of species occur, including high endemism in . Additional hotspots include , with notable diversity in and eastern , and the Pacific islands, where insular endemics such as variants of Pandanus tectorius thrive on remote atolls and high islands. Biogeographic patterns distinguish continental species, prevalent in African and Asian mainland forests, from insular forms dominant on oceanic islands; the latter often result from stepping-stone dispersal across into , facilitated by buoyant fruits dispersed via ocean currents, birds, and mammals. Some Pandanus species have been introduced beyond their native ranges through human activity, notably Pandanus utilis, originally from the in the , which has been transported to the , , southern , , and other Indian Ocean islands for its fiber-producing leaves. The genus generally occupies a envelope with optimal temperatures of 20–30°C and some tolerance for subtropical conditions, occurring from up to altitudes of 3,000 m, though most species are lowland dwellers below 1,000 m.

Habitat Preferences

Many species of Pandanus exhibit a strong preference for coastal and wetland environments, particularly in tropical and subtropical regions, where they colonize sandy beaches, mangrove fringes, riverbanks, and saline swamps. These habitats provide well-drained, sandy or peaty soils that support their prop-root systems, while tolerance for salt spray and periodic inundation allows establishment in brackish zones. For instance, Pandanus tectorius thrives on rocky and sandy shorelines near mangroves, adapting to high winds and direct sunlight through sunken stomata and thick cuticles that minimize water loss. Inland, Pandanus species occupy diverse microhabitats such as humid understories, freshwater swamps, and rocky outcrops in tropical lowlands and highlands. In New Guinea's highland s, species like Pandanus julianettii favor shaded slopes with moderate to heavy canopy cover (30–75%), occurring at altitudes from 1,300 to 3,300 m in areas with high (82–90%) and temperatures around 20–22°C. These inland niches often feature acidic clay soils rich in , contrasting with the alkaline preferences of coastal forms, though many species tolerate a broad edaphic range from well-drained sands to waterlogged peats with pH levels spanning acidic to alkaline (4.8–alkaline). Climatically, Pandanus dominates lowland humid but extends into montane rainforests and seasonally dry areas, with some xerophytic adaptations enabling survival in water-limited environments. Species like Pandanus forsteri store water in radicum tissues at prop-root tips, retaining up to 9.6 times their dry mass during droughts on isolated islands. Altitudinal niches vary, from sea-level coastal zones to up to 3,000 m in some populations, with overall preferences for high-rainfall regimes that support their growth in both wetlands and upland forests. In human-modified landscapes, Pandanus often persists in systems, such as rubber plantations and riverside clearings, where it benefits from partial shade and disturbed soils.

Ecology

Biotic Interactions

Pandanus exhibit a range of strategies, predominantly anemophily, where wind serves as the primary vector for transfer, as observed in . Certain , such as , display entomophilous traits, with thermogenic inflorescences attracting sap beetles (Amystrops spp.) as effective pollinators, evidenced by high loads on captured insects and exclusion experiments confirming reduced fruit set without beetle access. While biotic is less common across the genus, birds play a role in fruit dispersal. Herbivory on Pandanus primarily targets leaves and fruits, with browsers causing significant damage in and introduced ranges. Leaves, armed with sharp marginal spines, serve as a physical defense mechanism against folivores, deterring access by restricting movement and increasing handling time for herbivores, a strategy common in spiny monocots. Specific insect herbivores include planthoppers (e.g., Jamella australiae), which infest leaf sheaths and contribute to dieback, though direct leaf browsing by weevils remains poorly documented. Fruits are frequently consumed by vertebrates such as fruit bats ( spp.), rats (Rattus spp.), and land crabs (e.g., Coenobita spp.), acting as both predators that damage seeds and dispersers that facilitate propagation across islands. In dense stands, mammalian browsers like rats may also graze young leaves, exacerbating pressure on seedlings. Symbiotic relationships enhance Pandanus nutrient acquisition, particularly in nutrient-poor environments. Arbuscular mycorrhizal fungi (AMF) form associations with of like and Pandanus fascicularis, improving phosphorus and other mineral uptake in sandy or coastal soils, as confirmed by diversity and root colonization studies in rhizospheres. These symbioses are especially vital in oligotrophic habitats, where AMF colonization rates exceed 50% in natural populations. Evidence for nitrogen-fixing bacterial symbioses in of Pandanus is limited, though general suggests potential associations in flooded systems. Pathogens pose threats to Pandanus health, particularly in humid or introduced settings. Fungal leaf spots are prevalent, caused by species such as Nigrospora aurantiaca and Pestalotiopsis clavispora, which produce necrotic lesions on foliage of Pandanus amaryllifolius and other taxa, leading to reduced photosynthesis in affected stands. Viral diseases, including badnaviruses, induce mosaic and chlorotic symptoms in pandan grass (Pandanus amaryllifolius), as identified through high-throughput sequencing of symptomatic tissues. Invasive pests like the hala scale (Thysanococcus pandani) infest Pandanus tectorius in Pacific islands, sucking sap from leaves and causing yellowing, deformation, and dieback, with rapid spread via crawlers in non-native ranges such as Hawaii. Competitive interactions shape Pandanus distribution, with species demonstrating moderate that permits understory persistence in forested habitats. Photosynthetic acclimation in shaded leaves of species like Pandanus cookii maintains comparable rates to sun-exposed dicot trees, enabling survival under canopy cover. However, post-disturbance dynamics favor Pandanus as a pioneer in open, littoral, or burned areas, where fast growth allows establishment, though it may be outcompeted by more aggressive light-demanding species in prolonged successional gaps. In human-modified landscapes, with invasives further limits recruitment in niches.

Ecosystem Roles

Pandanus species fulfill critical provision roles in coastal and ecosystems, primarily through their extensive prop root systems that stabilize sandy dunes and riverbanks against . These anchor the plants in unstable substrates, such as those on small islands like in , where Pandanus populations have demonstrated effectiveness in maintaining sediment integrity and supporting adjacent vegetation communities. By creating elevated, branched structures, the prop roots form microhabitats that shelter epiphytes, birds, and ; for instance, in Australian coastal regions, they host specialist insects like the peppermint stick insect (Megacrania batesii), enhancing local faunal diversity. In nutrient cycling, Pandanus contributes substantially to soil enrichment in nutrient-poor environments. The decomposition of its nitrogen-rich leaf litter releases essential nutrients into sandy coastal soils, improving fertility and supporting plant succession in tropical settings. Wetland-adapted species, such as Pandanus amaryllifolius, further aid in water filtration by absorbing high levels of nitrates (up to 100% removal from 200 mg/L solutions over 14 days) and phosphates (64% removal from 100 mg/L over six weeks), thereby reducing eutrophication risks in aquatic systems. These plants also facilitate sediment trapping in hydrophytic conditions, promoting clearer water and stable substrates in tropical wetlands. Pandanus serves as a keystone species in insular and coastal ecosystems, bolstering biodiversity by providing food and habitat for specialist fauna. In Pacific island contexts, its fruits are dispersed by fruit bats such as flying foxes (Pteropus spp.), which consume the fleshy syncarps and excrete viable seeds, maintaining gene flow and forest regeneration across fragmented habitats. The plant's presence often indicates intact mangrove fringes, where it occupies swampy margins and supports associated invertebrate and avian communities, thereby signaling ecosystem health in tropical coastal zones. Regarding carbon sequestration, Pandanus exhibits moderate sequestration rates through aboveground and belowground biomass accumulation, with Pandanus tectorius storing lower but notable carbon levels compared to dominant forest trees in mixed coastal stands. Peat-forming wetland species enhance this capacity by contributing to long-term soil carbon storage, integrating into broader blue carbon dynamics of coastal systems where vegetation traps organic matter in sediments. Pandanus demonstrates resilience as a following disturbances such as cyclones and , rapidly colonizing exposed areas to facilitate . On vulnerable islands, dense patches of at least 4.53 m width can reduce heights, preserving habitat and aiding recovery in dynamic coastal environments. In low-lying areas, however, rising levels pose challenges to this role by inundating root zones, potentially shifting Pandanus distributions and altering succession patterns.

Human Uses and Conservation

Traditional and Cultural Uses

In Pacific Island cultures, particularly among and , Pandanus leaves have long been a primary material for mats, baskets, hats, and traditional garments due to their durable fibers. In , women weave fine pandanus leaves into ta'ovala, ceremonial waist mats worn by both men and women during formal occasions to signify respect and . Similarly, in Southeast Asian Austronesian communities like those in 's Island, pandanus mats called sinasa are crafted for ceremonial use and daily flooring, often dyed using natural pigments from plants such as and leaves to produce vibrant yellows, greens, and reds. Pandanus serves significant roles in traditional diets across and , where young fruits and shoots provide edible components after preparation to remove irritants like . In Papua New Guinea's highlands, the complex of Pandanus species yields nutritious fruits and nuts that are roasted or boiled, forming a staple for indigenous groups like the Wopkaimin, who distinguish edible varieties through detailed ethnobotanical knowledge. The leaves of , known as pandan, are widely used in to impart a fragrant, nutty flavor to dishes, desserts, and beverages, tied to ancient Austronesian culinary practices. Medicinal applications of Pandanus span Indian and Pacific traditions, with infusions from leaves and roots employed to treat ailments such as and skin conditions. In Ayurvedic practices in , Pandanus odoratissimus leaves are applied topically for , , and wounds, while root extracts address spasms and , supported by their and properties. Extracts from fragrant pandan leaves have also shown efficacy in accelerating incisional through ointments that promote tissue regeneration. In some Indian contexts, the plant's bark provides for dyeing and minor wound treatments, though its use is less documented in continental African traditions beyond . Pandanus holds sacred and symbolic significance in and Austronesian societies, often linked to creation stories, fertility, and spiritual protection. In Micronesian lore, such as among the Gilbertese, pandanus rituals invoke abundance for fruit crops and are integral to and ceremonies, reflecting its role as a life-sustaining . Samoan myths associate the Auriaria, patron of head-hunting rituals, with a primordial pandanus tree named "The Ancestress," symbolizing ancestral origins and ritual power. Across Austronesian groups in the Pacific, pandanus leaves feature in body adornments like wreaths and purification rituals, while their prop roots and leaves construct elements such as thatched roofs and walls, embodying communal and spiritual harmony. Historical trade in Pandanus materials facilitated cultural exchanges during Austronesian migrations and colonial eras, with leaves serving as key commodities. From , bundles of Pandanus utilis leaves, locally called vacoa, have been harvested and exported to neighboring islands like and for thatching roofs and weaving baskets, a practice rooted in pre-colonial resource networks. These migrations, originating from around 3000–1500 BCE, spread Pandanus cultivation and weaving techniques across the Pacific, enhancing inter-island trade in fibers and dyes.

Cultivation and Modern Applications

Pandanus species are primarily propagated vegetatively through suckers or offsets, which allows for rapid clonal and preservation of desirable traits such as fragrance or quality. Stem cuttings of 20-40 cm length, with trimmed leaves, can also be planted directly into well-drained , achieving high success rates in tropical conditions. Seed is possible but slower, as seeds often exhibit requiring soaking for several days before , with seedlings taking 4-12 months to reach transplant size; this method is less common due to variability in offspring. In temperate regions, cultivation under controlled humidity and temperature facilitates introductions of species like . Cultivation of Pandanus thrives in tropical climates with temperatures above °C, where require well-drained, sandy or loamy soils to prevent , often in coastal or alkaline conditions with 6-10. Full sun exposure of 6-8 hours daily promotes robust growth and fruiting, though partial shade protects young from excessive wind; spacing in commercial plantations typically ranges from 3-5 meters to accommodate mature heights up to meters and support development. is essential during dry seasons to maintain without waterlogging, with established showing tolerance to periodic but benefiting from supplemental water for optimal leaf and fiber production. Commercial products derived from Pandanus include essential oils extracted from leaves and flowers, particularly from P. odoratissimus, which are used in perfumes for their floral notes and in food flavorings to impart a vanilla-like aroma in desserts and beverages. In , variegated cultivars like P. veitchii (syn. P. tectorius 'Veitchii') are popular ornamentals for landscapes and indoor settings due to their striking white-margined leaves, grown in pots or as feature plants in warm climates. Industrial applications of Pandanus encompass leaf extracts incorporated into for their and aromatic properties, enhancing products like lotions and shampoos with natural scents and skin-soothing benefits. Sustainable from Pandanus leaves supports eco-tourism initiatives, where communities in Pacific islands produce handicrafts like mats and baskets using harvested s, promoting through renewable sourcing. Genetic improvement efforts, including breeding programs assessing diversity via markers like ISSR and SSR, aim to enhance quality for stronger, more durable materials in textiles and composites. Global trade in Pandanus products, such as dried leaves, extracts, and woven goods, is led by major producers and , with exporting over 100 shipments of pandan leaves and powders annually valued at approximately $125,000. Overharvesting poses challenges to wild populations, prompting adoption of sustainable certifications like those under guidelines for non-timber forest products to ensure ethical sourcing and long-term viability.

Conservation Status

Of the approximately 135 Pandanus species assessed by the , around 26% are classified as threatened with extinction (Critically Endangered, Endangered, or Vulnerable), while 26 species are categorized as , primarily owing to limited surveys in remote island ecosystems across the Pacific and Indian Oceans. This underassessment highlights the genus's vulnerability, as many of the estimated 700–800 total remain unevaluated, particularly endemics in isolated archipelagos where baseline data is scarce. The main threats to Pandanus species stem from driven by , agricultural conversion, and coastal development, which fragment coastal forests and swamps where the plants thrive. exacerbates these pressures through sea-level rise and erosion of coral atolls, reducing suitable habitats for strandline species in low-lying Pacific islands like . Additionally, overexploitation for leaves used in weaving and other traditional crafts has depleted populations in the region, where unsustainable harvesting outpaces natural regeneration. Particularly vulnerable are insular endemics, such as subpopulations of Pandanus tectorius (hala) in Hawaii, which face local declines from invasive pests like the Pandanus scale insect (Thysanococcus pandani) and habitat loss, despite the species' global Least Concern status. Other examples include Pandanus aldabrensis (Critically Endangered) on the Aldabra atoll, threatened by invasive species and habitat degradation, and Pandanus glaucifer (Endangered) in Madagascar, impacted by forest clearance. In Africa, coastal species like Pandanus candelabrum experience pressures from development, contributing to its Least Concern but regionally precarious status. Conservation efforts include in situ protection within national parks, such as Indonesia's , which safeguards diverse Pandanus habitats in Papua's lowland forests. Ex situ collections in botanic gardens, like those managed by the Indonesian Institute of Sciences, preserve genetic material for restoration, while community-based programs in Pacific nations promote sustainable harvesting techniques for weaving materials to reduce . For instance, initiatives in and Noosa (Australia) involve local groups in monitoring and replanting to bolster coastal resilience. Key research gaps persist, including the need for updated phylogenies to better identify evolutionary distinct lineages for priority conservation, as current taxonomic uncertainties hinder targeted actions in biodiversity hotspots like . Legal protections vary by region; while no Pandanus species are listed under Appendix II, national laws in Pacific island nations, such as Hawaii's regulations against invasive threats and Indonesia's acts, regulate harvesting and to curb exploitation.

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