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Casuarina
Casuarina
Casuarina
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Not to be confused with Casuarius.
For other uses, see Casuarina (disambiguation).

Casuarina
Casuarina equisetifolia, showing red female flowers and mature fruits
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Fagales
Family: Casuarinaceae
Genus: Casuarina
L.[1]
Type species
Casuarina equisetifolia[2]
L.
Species

See text

Female cones of C. equisetifolia

Casuarina, also known as she-oak, Australian pine[3][4] and native pine,[5] is a genus of flowering plants in the family Casuarinaceae, and is native to Australia, the Indian subcontinent, Southeast Asia, islands of the western Pacific Ocean, and eastern Africa.

Plants in the genus Casuarina are monoecious or dioecious trees with green, pendulous, photosynthetic branchlets, the leaves reduced to small scales arranged in whorls around the branchlets, the male and female flowers arranged in separate spikes, the fruit a cone containing grey or yellowish-brown winged seeds.

Description

[edit]

Plants in the genus Casuarina are dioecious trees (apart from C. equisetifolia that is monoecious), with fissured or scaly greyish-brown to black bark. They have soft, pendulous, green, photosynthetic branchlets, the leaves reduced to scale-like leaves arranged in whorls of 5 to 20 around the branchlets. The branchlets are segmented at each whorl with deep furrows that conceal the stomates. Male flowers are arranged along branchlets in spikes with persistent bracteoles, female flowers in spikes on short side-branches (effectively "peduncles") that differ in appearance from vegetative branchlets. After fertilisation, the female spikes develop into "cones" with thin, woody bracteoles that extend well beyond the cone body. The cones enclose grey or yellowish-brown winged seed known as samaras.[6][7][8][9][10][11]

Ecology

[edit]

Casuarina are attacked by a range of herbivorous insects.

Taxonomy

[edit]

The genus Casuarina was first formally described in 1759 by Carl Linnaeus in Amoenitates Academicae and the first species he described (the type species) was Casuarina equisetifolia.[2][17] The generic name is derived from the Malay word for the cassowary, kasuari, alluding to the similarities between the bird's feathers and the plant's foliage.[18]

Species List

[edit]

The following is a list of Casuarina species accepted by Plants of the World Online as of April 2023:[19]

In 1982, Lawrence Johnson raised the genera Allocasuarina and Gymnostoma in the Journal of the Adelaide Botanic Gardens, and transferred some species previously included in Casuarina to the new genera. The species of Allocasuarina previously in Casuarina are: A. acuaria, A. acutivalvis, A. campestris, A. corniculata, A. decaisneana, A. decussata, A. dielsiana, A. distyla, A. drummondiana, A. drummondiana, A. fraseriana, A. grevilleoides, A. helmsii, A. huegeliana, A. humilis, A. inophloia, A. lehmanniana subsp. lehmanniana, A. littoralis, A. luehmannii, A. microstachya, A. monilifera, A. muelleriana, A. nana, A. paludosa, A. paradoxa, A. pinaster, A. pusilla, A. ramosissima, A. rigida, A. robusta, A. striata, A. tessellata, A. thuyoides, A. torulosa, A. trichodon and A. verticillata. The species of Gymnostoma previously included in Casuarina are G. chamaecyparis, G. deplancheanum, G. intermedium, G. leucodon, G. nobile, G. nodiflorum, G. papuanum, G. poissonianum, G. rumphianum and G. sumatranum and G. webbianum.[20]

Invasive species

[edit]
Casuarina on Gold Rock Beach, Grand Bahama

C. cunninghamiana, C. glauca and C. equisetifolia have become naturalized in many countries, including Argentina, Bermuda, Cayman Islands, Cuba, China, Egypt, Israel, Iraq, Mauritius, Kenya, Mexico, Brazil, South Africa, the Bahamas,[21] and Uruguay. They are considered an invasive species[22][23] in the United States, especially in southern Florida[24] where they have nearly quadrupled in number between 1993 and 2005 and are called the Australian pine.[citation needed] C. equisetifolia is widespread in the Hawaiian Islands where it grows both on the seashore in dry, salty, calcareous soils and up in the mountains in high rainfall areas on volcanic soils.[citation needed] It is also an invasive plant in Bermuda, where it was introduced to replace the Juniperus bermudiana windbreaks killed by a scale insect in the 1940s.[25]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Casuarina

View on Grokipedia
from Grokipedia
Casuarina is a comprising approximately 17 species of trees and shrubs in the family , native to , the , , and islands of the western . These plants are characterized by their woody habit, with heights reaching up to 35 meters, and distinctive needle-like cladodes—photosynthetic branchlets that replace functional leaves, while true leaves are reduced to small, scale-like structures arranged in whorls of 5 to 20. Most species are dioecious, producing unisexual flowers that are wind-pollinated, leading to cone-like fruits containing winged samaras. The genus belongs to the order and is one of four genera in the family, alongside Allocasuarina, Ceuthostoma, and Gymnostoma, with the family totaling around 90 species overall. Fossil evidence indicates that Casuarina has origins dating back to the Eocene in . Species exhibit rough, fibrous bark that is typically fissured or scaly and gray-brown to black in color, with branchlet teeth numbering 6 to 20 per whorl, varying by species in ridge width, hairiness, and other traits. They are either monoecious or dioecious, except for C. equisetifolia, which can be monoecious. Ecologically, Casuarina species play significant roles in their native habitats, particularly in stabilizing soils in semiarid and coastal environments through their root systems and nitrogen-fixing symbiosis with the actinomycete , which enables them to thrive in nutrient-poor soils; they also form mycorrhizal associations. However, several species, such as C. equisetifolia, C. cunninghamiana, and C. glauca, have become invasive in non-native regions like and other subtropical areas, outcompeting local flora due to their rapid growth and allelopathic effects. Notable species include C. equisetifolia (commonly known as Australian or ), valued for its timber but problematic as a , and C. cunninghamiana, often found along riverbanks and capable of hybridizing with C. glauca. Economically, Casuarina trees are widely utilized for fuelwood, timber, and pulp production, particularly C. equisetifolia in tropical plantations; they are also planted for , windbreaks, and shade in , though their invasiveness limits such uses in sensitive ecosystems. Their fast growth and make them suitable for reclamation projects in degraded lands, contributing to soil improvement via .

Description and morphology

Physical characteristics

Casuarina species are evergreen trees that typically reach heights of 6 to 35 , characterized by slender, drooping branchlets that give the an equisetoid, or horsetail-like, appearance. The bark is rough, fibrous, and ranges from grey-brown to reddish-brown, often exfoliating in longitudinal strips. These trees exhibit secondary thickening in their persistent branches, contributing to their . Most Casuarina species are dioecious, bearing unisexual flowers on separate male and female plants, though C. equisetifolia is an exception as a monoecious . The leaves are highly reduced to scale-like structures, white or brown in color, fused laterally at their bases to form toothed sheaths arranged in whorls of 4 to 16 around the nodes of the branchlets. These branchlets are needle-like cladodes, ribbed and furrowed, with internodes that vary in length and serve as the primary photosynthetic organs, as the true leaves are non-photosynthetic and often shed soon after formation. The branchlets feature deep furrows alternating with ridges, containing vertically aligned rows of stomata (typically 2–3 per groove) for , along with epidermal hairs in some that aid in adaptation to environmental stresses. Reproductive structures include woody, cone-like infructescences that are pedunculate and pubescent when immature, housing winged samaras as seeds. Additionally, Casuarina roots develop nodules in symbiosis with actinomycetes, enabling and supporting growth in nutrient-poor soils.

Reproduction

Casuarina species exhibit dioecious or monoecious , with separate and individuals predominant in many taxa, though some species display both conditions on a single . cones, typically arranged in whorls along branchlets, produce abundant , while cones, also whorled, feature reduced, scale-like bracts that house ovules and develop into woody infructescences upon fertilization. Pollination is anemophilous, relying on wind dispersal of lightweight, dry grains from male cones, which can travel considerable distances to reach strobili. This occurs year-round in tropical species like , with peak flowering varying by locality from June to December; pollen viability remains high, often exceeding 90%, facilitating effective cross-pollination in open stands. Following , female cones mature into infructescences over 18-20 weeks, releasing numerous winged samaras—small, indehiscent with papery wings—that enable or dispersal. Seed production is prolific, with good crops forming annually after age 5 in many , yielding approximately 650,000–760,000 per kilogram in C. equisetifolia; samara viability persists for 1-5 years under cool, dry storage conditions at 3-16% moisture and temperatures of -7°C to 3°C. Germination of Casuarina seeds is , typically occurring within 10-14 days under warm, moist conditions in well-drained, light soils, with rates ranging from 20-80% depending on freshness and pretreatment. For with hard seed coats, such as C. equisetifolia, using concentrated enhances by breaking , often achieving rates above 50% when combined with soaking or treatments like 0.1 mM IAA. Successful establishment frequently involves symbiotic associations with ectomycorrhizal fungi, which improve nutrient uptake, particularly , in nutrient-poor soils during early seedling growth. While primarily sexual, Casuarina exhibits rare through natural vegetative propagation in disturbed contexts, such as root suckering or from basal shoots following damage, allowing clonal spread without seed involvement. has not been documented in the .

Taxonomy and phylogeny

Etymology and history

The name Casuarina was established by in 1759 in his work Amoenitates Academicae, derived from the Latin casuarius (), reflecting the resemblance of the plant's pendulous, needle-like branchlets to the feathers of the ; this traces back to the Malay word kesuari for the . Linnaeus's initial description was based on C. equisetifolia as the , which he named from specimens collected in the , marking the genus's entry into formal botanical classification. Over the subsequent centuries, the of Casuarina underwent significant revisions as more were described and relationships within the Casuarinaceae family were clarified. In the late , Australian botanist Lawrence Alexander Sidney Johnson restructured the family, separating several into new genera to better reflect morphological and anatomical differences; notably, in 1982, he established for with reduced, unisexual flowers and indehiscent cones, while Gymnostoma was recognized earlier in for more primitive, bisexual-flowered taxa from and the southwest Pacific, and Ceuthostoma for a small group with distinct cone valves. These changes reduced the circumscription of Casuarina proper to primarily distributed in , , and the Pacific, emphasizing dioecious reproduction and dehiscent cones as defining traits. Molecular phylogenetic studies in the early , utilizing genes such as matK and rbcL, have confirmed the of and the distinct genera within it, placing the family firmly in the order among core while highlighting its unique combination of reduced leaves, wind-pollinated flowers, and symbiotic as evolutionarily derived features. Recent taxonomic syntheses, including the database updated through 2023, recognize 14 accepted species in Casuarina, reflecting ongoing refinements based on integrated morphological and genetic data without further major generic splits.

Species diversity

The genus Casuarina comprises 14 accepted species according to current taxonomic assessments, with the majority endemic to various regions of and one extending to and the Pacific. These species are trees or shrubs adapted to diverse Australian environments, from coastal dunes to inland woodlands, and are distinguished by variations in branchlet , morphology, and preferences. Among the most prominent species is C. equisetifolia, a fast-growing tree reaching up to 35 m in height, native to coastal areas from India through Southeast Asia, northern Australia, and Pacific islands, where it thrives in sandy, saline soils and serves as a pioneer in dune stabilization. C. cunninghamiana, a tall riverine species up to 30 m, is endemic to eastern Australia and southeastern New Guinea, favoring stream banks and floodplains with its pendulous branchlets and tolerance for periodic inundation. C. glauca exhibits high salt tolerance, growing as a medium-sized tree (up to 15 m) in swampy estuarine fringes and mangrove margins along Australia's east coast, often forming dense stands in brackish conditions. In contrast, C. pauper is a smaller tree (up to 10 m) adapted to semi-arid inland zones of southern and eastern Australia, occurring in open woodlands on calcareous soils with low rainfall. Phylogenetic analyses using matK gene sequences confirm Casuarina as monophyletic within , with species clustering into clades reflecting geographic and ecological divergence, such as coastal versus inland forms. Infrageneric groupings have been proposed based on these phylogenies and morphological traits like valve number in cones and branchlet features, distinguishing broader alliances akin to historical sections Casuarina (equisetifolia-like, with 6-8 teeth per whorl) and Acicularia (cunninghamiana-like, with 4-5 teeth). Significant taxonomic revisions occurred in 1982, when L.A.S. Johnson segregated numerous species from the broader Casuarina sensu lato into the new genus Allocasuarina, based on differences in seed wing structure, chromosome numbers, and inflorescence types, reducing Casuarina to its current circumscription of about 15 species (now refined to 14). This reassignment resolved many synonyms, such as former Casuarina taxa like C. distyla and C. muelleriana now in Allocasuarina, while retaining core Casuarina species with indehiscent, winged seeds. Conservation concerns affect some species; for instance, C. cristata (belah), a woodland tree of inland eastern Australia, faces threats from land clearing, with associated ecosystems listed as endangered in parts of its range.

Distribution and habitat

Native range

The genus Casuarina is predominantly native to , where it exhibits a broad distribution spanning from tropical regions in northern and the to temperate areas in Victoria and southwestern . This range encompasses diverse climatic zones, with species adapted to both arid interiors and coastal environments across states including , , and . Outside Australia, the native range extends to , including regions such as , , , , and the , as well as the Pacific islands like , , , and the . Notably, C. equisetifolia has the widest distribution among the species, occurring naturally from and through to and various Pacific archipelagos. The genus comprises approximately 14 accepted species, with the majority—over 90%—endemic to , reflecting high levels of regional . Casuarina species typically inhabit coastal dunes, riverbanks, heathlands, and fringes, showing strong adaptations to challenging conditions such as sandy, saline, or nutrient-poor soils. These habitats often feature well-drained substrates near water bodies, where the plants' and salt tolerance enable persistence in exposed, unstable environments. Biogeographic patterns include disjunct distributions across island chains and , consistent with historical dispersal via ocean currents and ancient land connections. Fossil evidence supports Gondwanan origins for the family, with pollen records of casuarinacean affinity dating back to the and becoming prominent in the across , , and even . These ancient distributions indicate that ancestral Casuarina-like lineages evolved in southern floras before modern speciation and range fragmentation.

Introduced ranges

Casuarina species have been widely introduced outside their native ranges since the , primarily for , windbreaks, and . These introductions were facilitated by colonial plantings, government-sponsored forestry programs, and ornamental horticultural trade, leading to establishment in diverse regions across the globe. By the early , species such as C. equisetifolia and C. cunninghamiana were being propagated from seeds collected in and , with deliberate plantings documented in locations like and as early as the 1820s and 1860s, respectively. The has successfully established in tropical and subtropical climates worldwide, with ten of its 14 now present beyond their native distributions in approximately 150 countries. In , introductions date to the late , with C. equisetifolia and C. cunninghamiana planted extensively in for dune stabilization and , where they now occur across more than 20 countries including and . Early plantings in the included (1890s), while saw introductions pre-1895, and primarily after 1950; these were often as coastal barriers and ornamentals, resulting in naturalized populations along shorelines and riverine areas. In , large-scale initiatives in and since the mid-20th century have promoted C. equisetifolia for timber and fuelwood, while in , Mediterranean regions such as and the host introduced stands for similar ecological roles. Hybrids have formed in some introduced areas, notably in , where interspecific crosses between C. equisetifolia, C. glauca, and C. cunninghamiana exhibit intermediate morphologies and enhanced adaptability, complicating local management efforts. These hybrids, confirmed through molecular analysis, occupy coastal and disturbed habitats, contributing to the genus's persistence in subtropical environments. Currently, Casuarina plantations span millions of hectares globally, with alone accounting for approximately 500,000 hectares dedicated primarily to C. equisetifolia for industrial and rural uses (recent estimates post-2010). Extensive plantings also cover significant areas in , , and parts of , supporting objectives across more than 100 countries. It is reported as invasive and naturalized in Caribbean island ecosystems including , the , and the Bahamas (data from early 2000s cited in 2023 compendium), posing risks to coastal amid climate-driven habitat shifts.

Ecology

Ecological interactions

Casuarina species experience significant herbivory from , including psyllids in the Casuarinicola that feed on branchlets and foliage, as well as weevils such as Artipus floridanus that damage stems and roots. Browsing by mammals, including and wallabies in native Australian habitats, targets young shoots and foliage, though such pressure is moderated by the plant's production of chemical defenses like condensed , which deter feeding by reducing and digestibility. Seeds of Casuarina are subject to predation and dispersal by birds, particularly cockatoos such as the glossy black-cockatoo (Calyptorhynchus lathami), which extract and consume the majority of viable seeds from cones, often leaving little for but occasionally dispersing uneaten seeds through droppings. In native ranges, also play a in seed predation and dispersal, harvesting seeds for food stores but sometimes abandoning them intact in nest middens where they can sprout. Casuarina trees exert competitive effects on understory flora through their dense canopies, which limit light penetration, and via , where leachates from leaves and litter release inhibitory phenolics that suppress and growth of neighboring . In fire-prone ecosystems, Casuarina species are well-adapted to frequent bushfires, resprouting vigorously from woody lignotubers that store carbohydrates and enable rapid regeneration of shoots post-fire, thereby maintaining dominance in disturbed habitats. Casuarina species are also susceptible to fungal pathogens like Fusarium spp., which can affect root health and in native habitats. Beyond their well-known nitrogen-fixing , Casuarina forms mutualistic associations with arbuscular mycorrhizal fungi, which enhance nutrient uptake, particularly , in nutrient-poor soils and improve tolerance to environmental stresses like . is primarily anemophilous, relying on dispersal of .

Nitrogen fixation and soil role

Casuarina species form symbiotic associations with the actinomycete genus in specialized root nodules, where the convert atmospheric (N₂) into , providing a key source of fixed for the plant. This actinorhizal symbiosis enables Casuarina to thrive in nitrogen-poor environments, as colonizes the roots and facilitates through the expression of enzymes within the nodules. Through this process, Casuarina significantly enhances , particularly in nutrient-deficient sandy substrates, by increasing via leaf litter decomposition and elevating over time. In coastal dunes and other infertile sites, the fixed enriches the profile, supporting greater microbial activity and nutrient cycling. Fixation rates can reach up to 200 kg N/ha/year in settings, though typical values range from 40 to 94 kg N/ha/year depending on environmental conditions and . As a , Casuarina plays a crucial role in by stabilizing shifting sands in dunes through its extensive systems and dense foliage, which trap sediments and reduce . This initial improves site conditions, allowing subsequent establishment of less tolerant plant species and fostering in early successional stages. Nitrogen fixation efficiency varies among Casuarina species, with C. glauca demonstrating particularly high performance in saline soils due to its tolerance of high salt levels, which minimally disrupts the Frankia symbiosis compared to other species like C. equisetifolia. This adaptation allows C. glauca to maintain effective nitrogen input in coastal or brackish environments where soil salinity inhibits many other plants.

Human interactions

Cultivation and uses

Casuarina species are propagated primarily through seeds or stem cuttings to facilitate widespread planting in managed systems. , which number approximately 300,000 per pound, germinate readily in 4-8 days without pretreatment, though soaking in warm for several hours enhances viability; they are sown in well-draining media and kept moist until establishment. Vegetative propagation via semi-hardwood cuttings taken in late spring or summer, treated with rooting hormone, and placed in a moist sandy medium under high achieves rooting in several weeks, enabling clonal production of superior genotypes. These trees exhibit strong tolerance to once established, requiring supplemental only during the first few months to develop deep roots, and to , thriving in coastal soils exposed to salt spray. For plantings, spacing of 2-3 meters between trees optimizes and reduction while allowing growth; wider intervals of 3-4 meters suit medium-sized species in rows. In landscaping, Casuarina trees are valued for their feathery, needle-like branchlets that provide an aesthetic, pine-like appearance, and their extensive root systems contribute to erosion control by stabilizing sandy or coastal soils. The wood of Casuarina is a dense hardwood suitable for furniture, construction poles, and tool handles due to its strength and termite resistance, while its high calorific value makes it a preferred source for fuelwood and charcoal production. In tropical regions, these fast-growing species achieve annual height increments of up to 2 meters, enabling harvests within 7-10 years for pulpwood or poles. Beyond timber, Casuarina serves multiple roles in and rural economies; its branchlets provide low-nutritive but emergency for such as and , supporting in arid conditions. The trees are integrated into systems to enhance via and are commonly planted as windbreaks in coastal and agricultural areas to protect crops from salt-laden winds and reduce erosion. Casuarina has been planted extensively for economic purposes since the 1800s, with introductions to India in the 19th century for fuelwood and coastal stabilization, and to regions in Africa such as Kenya for similar agroforestry applications.

Invasiveness

Several species within the genus Casuarina have established as invasive exotics in non-native regions, often forming dense monocultures that dominate landscapes and suppress indigenous flora. Casuarina equisetifolia is particularly problematic in coastal areas of Florida and Hawaii, USA, where it rapidly colonizes beaches and dunes, outcompeting native species and altering habitat structure. Similarly, C. glauca invades riparian and coastal zones in Florida, USA, while C. cunninghamiana threatens riverine habitats in Florida, USA, by forming thickets that exclude local vegetation. These invasions are exacerbated by the trees' fast growth rates, reaching heights of up to 10 meters in a few years, and their ability to thrive in nutrient-poor, sandy soils. The ecological impacts of invasive Casuarina species are profound, primarily through competition with native plants via allelopathy, shading, and resource monopolization. Rapid establishment leads to the displacement of understory species, drastically reducing biodiversity in affected areas; for example, invaded coastal dunes often exhibit near-total loss of native herbaceous cover and diminished shrub layers. These trees also alter hydrology by stabilizing soils and increasing evapotranspiration, which can lower groundwater levels and disrupt wetland dynamics in sensitive ecosystems. Fauna suffer indirectly, as the simplified monocultures provide fewer food sources and nesting sites, contributing to declines in pollinators, birds, and reptiles dependent on diverse native communities. Spread of invasive Casuarina occurs primarily through wind-dispersed samaras, which can travel long distances along coastlines, aided by currents and storms; human-mediated via landscaping debris and vehicles further accelerates establishment in disturbed sites like roadsides and cleared lands. Seeds germinate readily in bare or sandy substrates, enabling pioneer colonization that prevents native succession. Management of Casuarina invasions relies on integrated approaches, including mechanical removal of seedlings and saplings by hand-pulling or cutting, which is effective for small-scale infestations but labor-intensive for larger stands. Herbicides such as applied to cut stumps or via foliar spray provide control for mature trees, often requiring follow-up treatments to address resprouting, with success rates improving when combined with monitoring. Biological control efforts are promising, with host-specific agents like the Asian seed-feeding wasp Bootanelleus orientalis undergoing trials in the 2020s to reduce seed viability and limit spread in regions like . Regulatory measures reflect the severity of these invasions: C. equisetifolia and related species were designated as noxious weeds in , , during the 1990s under state prohibited plant lists, banning sale, transport, and propagation. In , Casuarina species are scheduled as invasive plants requiring eradication or control on public lands. Globally, invasions are tracked through databases like the IUCN Global Invasive Species Database and CABI Compendium, with updates as recent as 2023 highlighting ongoing risks and distribution patterns.

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  60. https://ecocrop.apps.fao.org/ecocrop/srv/en/cropView?id=646
  61. https://www.srs.fs.usda.gov/pubs/misc/ag_654/volume_2/casuarina/casaurina.htm
  62. https://pfaf.org/user/Plant.aspx?LatinName=Casuarina%2Bequisetifolia
  63. https://apps.worldagroforestry.org/treedb/AFTPDFS/Casuarina_cunninghamiana.PDF
  64. https://data.fs.usda.gov/research/pubs/iitf/sm_iitf056%20%20(11).pdf
  65. https://www.cal-ipc.org/plants/risk/casuarina-equisetifolia-risk/
  66. https://www.ars.usda.gov/ARSUserFiles/33272/Gaskin_243079.pdf
  67. https://edis.ifas.ufl.edu/publication/WG209
  68. https://tsusinvasives.org/home/database/casuarina-equisetifolia
  69. https://www.fdacs.gov/Agriculture-Industry/Pests-and-Diseases/Plant-Pests-and-Diseases/Noxious-Weeds/Casuarina-spp.-Australian-Pines
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