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Liquidambar

Liquidambar is a of flowering trees in the family , comprising approximately 15 of large and trees native to eastern , eastern , and the eastern . These trees are characterized by their alternate, long-stalked leaves—often star-shaped and lobed in temperate species, with unlobed forms in subtropical ones—and their monoecious flowers arranged in yellowish-green globular heads, maturing into distinctive woody, spiny infructescences. The genus name, derived from the Latin liquidus (liquid) and the anbar (), alludes to the fragrant, -like produced by the , which has been historically used in perfumes, medicines, and adhesives. Notable species include , the American sweetgum, which is widespread in the and valued for its vibrant autumn foliage in shades of red, purple, and yellow, as well as its commercial timber used in furniture, , and . In , and Liquidambar acalycina are prominent, with the former native to , , , and parts of , featuring similar deciduous habits and ecological roles in mixed forests. , found in southwestern and nearby , is an evergreen species prized for its storax resin, a key ingredient in ancient and modern fragrances. Members of the typically grow to 20–40 meters in height, with fissured and ovoid buds, thriving in moist, well-drained soils but tolerant of periodic flooding; however, their brittle wood makes them susceptible to storm damage. Ecologically, Liquidambar support diverse , providing and food, while their seeds—often winged but with low fertility rates—aid in dispersal. Cultivated widely for ornamental purposes due to their striking fall colors and shade, they are also significant in , with L. contributing to efforts in the U.S. South.

Taxonomy

Etymology

The genus name Liquidambar derives from the Latin word liquidus, meaning "liquid," and the ambar, meaning "amber," alluding to the fragrant, resinous sap exuded by the tree that resembles liquid in appearance and aroma. This nomenclature was established by the Swedish botanist in his in 1753, where he described the based on the distinctive properties of the exudate from species like . The common name "sweetgum" originates from the sweet-tasting historically harvested and chewed as a by Native American communities and early European settlers in . Regional variations include "storax," derived from the term styrakos for a fragrant , reflecting the tree's liquid known as American storax. The linguistic roots of these names trace back to ancient Mediterranean uses of similar resins in perfumery and ; and Romans employed storax-like balsams for , ointments, and as a in wine, as documented by classical authors such as .

Extant species

The genus Liquidambar comprises 15 accepted extant species, as recognized in a comprehensive taxonomic synopsis of the family . These species exhibit disjunct distributions, with the majority occurring in (particularly , , and the eastern Mediterranean), one in eastern extending to , and none in or . The genus was recently reclassified within the segregate family , distinct from the broader Hamamelidaceae, based on molecular and morphological evidence that supports the inclusion of former genera Altingia and Semiliquidambar under Liquidambar. Species distinctions often involve leaf lobing (ranging from 3 to 7 lobes, with some showing polymorphic variation), tree height (typically 20–40 m, though smaller in some), and fruit characteristics (woody capsules 2–4 cm in diameter, varying in spine length and density). Several species have synonyms from prior generic placements, such as Altingia cambodiana for L. cambodiana. Key species include L., the American sweetgum, native to eastern from to and west to , extending southward to (, , , ). This large reaches up to 40 m tall with a pyramidal to rounded crown, featuring alternate, star-shaped leaves (7–19 cm long, 5–16 cm wide) typically with 5–7 lobes and rounded-toothed margins, shiny dark green above and turning brilliant red, purple, or in autumn. Fruits are spiny, globose capsules (2.5–4 cm diameter). Mill., the Oriental sweetgum, is endemic to the eastern region, including southwestern , (), and , where it forms pure stands in floodplains. It is a smaller (15–30 m tall) with coarsely toothed, 5-lobed leaves (to 8 cm across), each lobe further divided, and unlobed or shallowly lobed forms common; fall color ranges from to purple. Fruits are smaller (about 2 cm) with shorter spines compared to L. styraciflua. Hance, the Formosan sweetgum, is distributed across , including southern , , central and southern , and . This grows 30–40 m tall with a pyramidal to rounded crown, distinguished by 3-lobed (rarely 5-lobed) star-shaped leaves (10–15 cm wide) with serrated margins and acuminate tips, often pubescent on young shoots. Fruits are globose, 3–4 cm in diameter, with persistent styles. Other notable species are predominantly Asian and often rarer, with several exhibiting polymorphic leaf shapes (varying from unlobed to multi-lobed within individuals). Liquidambar acalycina H.T. Chang (synonym Semiliquidambar acalycina), native to southern , is a slender to 25 m with 3- (rarely 5-) lobed leaves (5–10 cm long), glabrous and turning red in fall; it is considered rare in . Liquidambar cambodiana (Lecomte) Ickert-Bond & J.Wen (synonym Altingia cambodiana), restricted to , is a tropical with shallowly 3–5-lobed leaves and small fruits (2 cm), known from few collections. Liquidambar caudata (H.T. Chang) Ickert-Bond & J.Wen (synonym Semiliquidambar caudata), from southern , shows highly variable leaf morphology (3–7 lobes, sometimes unlobed) and reaches 20–30 m; fruits have prominent persistent styles. The remaining species, such as L. chingii (southern to , 3–5-lobed leaves with suprabasal venation), L. chinensis (southern to , , 5-lobed leaves), L. excelsa ( to , large 5–7-lobed leaves), L. gracilipes (, slender leaves), L. multinervis (northern , , 3–5-lobed leaves with strong venation), L. nobilis (, multi-lobed polymorphic leaves), L. obovata (, , shallowly lobed leaves), L. poilanei (, rare with 3-lobed leaves), L. siamensis (southern to , 5-lobed leaves), and L. yunnanensis (southeastern , , to , variable lobing), further illustrate the genus's diversity in subtropical to tropical forests, with many sharing resinous properties but differing in fruit spine density and leaf pubescence.
SpeciesNative RegionDistinguishing Traits
L. acalycina H.T. ChangSouthern ChinaSlender tree to 25 m; 3–5-lobed glabrous leaves (5–10 cm), red in fall; small fruits (2–3 cm).
L. cambodiana (Lecomte) Ickert-Bond & J.WenCambodiaTree to 20 m; shallowly 3–5-lobed leaves; compact spiny fruits (2 cm); rare.
L. caudata (H.T. Chang) Ickert-Bond & J.WenSouthern China20–30 m tall; polymorphic leaves (3–7 lobes or unlobed); fruits with long styles.
L. chingii (F.P.Metcalf) Ickert-Bond & J.WenSouthern China to VietnamTree to 25 m; 3–5-lobed leaves with unique venation; smaller stature.
L. chinensis Champ. ex Benth.Southern China to Vietnam20–30 m; evergreen, 5-lobed leaves; fruits 3 cm with dense spines.
L. excelsa (Noronha) OkenBhutan, China (SE Yunnan), MalesiaLarge tree to 40 m; 5–7-lobed broad leaves (15 cm); tropical fruits 4 cm.
L. formosana HanceS. Korea, Taiwan, C/S China, N. Vietnam30–40 m; 3-lobed star leaves (10–15 cm), pubescent petioles; globose fruits 3–4 cm.
L. gracilipes (H.T. Chang) Ickert-Bond & J.WenChinaSlender tree; narrow 3–5-lobed leaves; rare, with elongated fruits.
L. multinervis (W.C.Cheng) Ickert-Bond & J.WenChina (N. Guizhou)Tree to 20 m; 3–5-lobed leaves with prominent venation; fruits 2–3 cm.
L. nobilis (H.T. Chang) Ickert-Bond & J.WenChina20 m; polymorphic multi-lobed leaves; compact crown.
L. obovata (Merr. & Chun) Ickert-Bond & J.WenChina (Hainan)Tree to 20 m; shallowly 3–5-lobed or unlobed leaves; small spiny fruits (~2 cm); rare.
L. orientalis Mill.E. Mediterranean (Turkey, Greece, Cyprus)15–30 m; 5-lobed leaves (8 cm) with secondary lobing; smaller spiny fruits (2 cm).
L. poilanei (Tardieu) Ickert-Bond & J.WenVietnamTree to 25 m; 3-lobed leaves; endemic and rare.
L. siamensis (Craib) Ickert-Bond & J.WenS. China to Thailand, Vietnam30 m; 5-lobed leaves; fruits with medium spines (3 cm).
L. styraciflua L.E. North America to Central America40 m; 5–7-lobed star leaves (7–19 cm), fall color; spiny fruits 2.5–4 cm.
L. yunnanensis (Rehder & E.H. Wilson) Ickert-Bond & J.WenSE China (Yunnan) to Vietnam25–35 m; variable 3–5-lobed leaves; polymorphic forms common.

Fossil record

The genus Liquidambar originated in the , with the earliest known s consisting of leaf impressions from n deposits dating to approximately 80 million years ago. inflorescences assigned to the genus also appear in Upper Cretaceous strata of , marking the initial diversification of . The genus achieved peak diversity during the period ( to ), with numerous species documented across Laurasian continents through leaves, fruits, and . Key fossil species illustrate this historical range and morphological variation. Liquidambar changii, from the Middle of central , , is represented by anatomically preserved infructescences that reveal details of fruit structure and development, including 1–2 mature seeds per carpel alongside aborted ovules. In contrast, Liquidambar maomingensis from the late Eocene of the Basin, , features polymorphic leaves with 3–5 lobes and associated capitate infructescences, suggesting an ancestral polymorphic lineage within the . Other notable taxa span the Eocene to , such as Liquidambar bella from the Eocene of Hainan Island, China, and various leaf forms from European sites. A recent 2024 discovery of structurally preserved infructescences, leaves, and from the late Nanning Basin, , provides new evidence of mechanisms through winged structures and highlights transitional morphologies between Liquidambar and related altingioid genera. During the Tertiary, exhibited a broad distribution across the , including , , and , reflecting warm, mesic paleoenvironments. Pleistocene glaciation restricted its range, resulting in current relict populations and local extinctions, particularly from and western . Evolutionarily, the genus's disjunct modern distribution—spanning eastern , western , and eastern —arises from vicariance following the divergence of lineages in the early , as supported by chloroplast DNA analyses.

Biology

Morphology

Liquidambar are large trees, primarily in temperate regions but some subtropical are , typically reaching 20–40 m in though sizes vary by , with straight trunks and crowns that transition from pyramidal in youth to rounded and spreading in maturity. They exhibit a monoecious habit, with reproductive structures on the same . Mature individuals often develop buttressed bases at the trunk for enhanced stability. Leaves are alternate and long-stalked. In species, they are palmately lobed with 3–7 lobes, up to 20 cm long, giving them a distinctive star-shaped appearance with serrated margins and long petioles; they are glossy green during the but renowned for their vibrant autumn coloration in , , and , resulting from the accumulation of anthocyanins in response to environmental cues such as cooler temperatures and increased . species have unlobed, entire or serrated leaves that remain green year-round. Stems and branches feature grayish that becomes deeply furrowed with age, often developing corky ridges on younger branches for added . The is notable for resin canals distributed throughout the stems, leaves, and other vegetative tissues, which produce aromatic, ous exudates upon injury. Growth rates are moderate to fast, particularly on moist, well-drained sites, allowing the trees to disturbed areas and form dense stands.

Reproduction

Liquidambar are monoecious, bearing separate flowers on the same tree within separate inflorescences. Male flowers form pendulous catkins consisting of racemes of staminate florets, while female flowers develop in upright, globose spikes or heads of pistillate florets; both are small, greenish, and non-showy. These wind-pollinated flowers typically bloom from March to early May, varying by and . Following , female inflorescences mature into aggregate fruits composed of numerous woody capsules that form distinctive spiny, spherical structures commonly known as "gumballs," measuring 2–4 cm in diameter. These fruits persist on the through winter, turning dark brown and remaining intact until dehiscing in late winter or early spring to release seeds. Each fruit contains up to 100 small , typically 1–2 per individual capsule within the aggregate, with thin wings facilitating anemochory or dispersal. occurs primarily in autumn and winter, with most seeds falling within 60 m of the parent tree, though distances up to 180 m have been recorded. Flowering in Liquidambar is phenologically synchronized within populations, often producing fair crops annually and abundant crops every few years, with trees capable of from 20–30 years of age onward. The genus's production, including in floral and foliar tissues, plays a role in deterring herbivores during vulnerable reproductive phases by chemically inhibiting feeding.

Distribution and ecology

Geographic distribution

The genus Liquidambar exhibits a disjunct native distribution across three primary regions: eastern , the , and East and . In eastern North America, L. styraciflua is the sole species, ranging from southern and southern southward through the to central and eastern , extending westward to , , and , and further into northeastern and montane regions of as far south as . In the eastern Mediterranean, L. orientalis is endemic to southwestern and the Greek island of , occurring primarily in and habitats up to 1,800 meters . The majority of species are concentrated in East and Southeast Asia, including L. formosana in central and southern , , , and ; L. chinensis in southern China to ; L. excelsa from through southeastern , China, to western ; L. cambodiana in ; and L. acalycina in southern China. These Asian species typically inhabit subtropical and temperate forests. Beyond their native ranges, Liquidambar species have been widely introduced and cultivated as ornamentals in temperate and subtropical regions worldwide. L. styraciflua and L. formosana are commonly planted in , including the and , as well as in , , and , where they thrive in urban and landscaped settings. In some introduced areas, such as , L. styraciflua has naturalized in disturbed sites. This disjunct biogeographic pattern reflects a relict distribution from the period, shaped by , the fragmentation of the Tethys seaway, and subsequent climate oscillations that isolated populations across continents. For instance, L. styraciflua occupies lowland coastal plains and rich bottomlands in the southeastern U.S., while Asian species like L. formosana are found in moist subtropical forests of and .

Habitat and ecological role

Liquidambar species thrive in a variety of temperate to subtropical environments, predominantly favoring moist, well-drained soils such as alluvial clays and loams with medium and imperfect drainage. They are commonly found in bottomlands, floodplains, river valleys, and moist woodlands, where they tolerate periodic flooding but often prefer upland sites with consistent moisture availability. preferences range from slightly acidic to neutral, though some like adapt to alkaline conditions in hydromorphic soils rich in and low in . Annual rainfall in their habitats typically exceeds 1000 mm, supporting their growth in climates with 180–320 frost-free days and temperatures ranging from -21°C to 38°C. Ecologically, Liquidambar acts as a in disturbed habitats, readily colonizing old fields, logged areas, and post-fire sites to form dense, nearly pure stands that facilitate in upland and ecosystems. Their spiny, ball-like fruits provide a vital source for , including birds, squirrels, and chipmunks, while the trees offer shelter and nesting sites, enhancing in mixed forests. The resin exuded by Liquidambar species serves as an defense mechanism, protecting against pathogens and wounds, which contributes to their resilience in dynamic environments. Additionally, the colorful fall foliage decomposes to enrich , playing a key role in nutrient cycling by returning essential elements like and carbon to the . Liquidambar engages in several interactions that bolster its ecological integration. Primarily wind-pollinated, the trees occasionally attract to their catkins, supporting communities indirectly. They form symbiotic associations with vesicular-arbuscular mycorrhizal fungi, such as Glomus mosseae, which enhance nutrient uptake—particularly —and improve establishment in nutrient-poor soils. These mutualisms underscore Liquidambar's contribution to and stability. While the genus faces no global conservation threats, disjunct populations are vulnerable to , which may alter moisture regimes and temperature extremes, potentially contracting suitable habitats for species like L. orientalis in the Mediterranean. Local rarity persists in relict stands, such as those of L. orientalis in , where habitat loss has reduced forest area from 7000 ha to 1657 ha over two centuries due to pressures. In , broader distributions remain stable.

Uses and cultivation

Ornamental and medicinal uses

Liquidambar species, particularly , are widely appreciated for their ornamental value in and . These trees are planted for their vibrant fall foliage, which transforms into shades of yellow, orange, red, and purple, creating striking seasonal displays that enhance parks, streets, and residential areas. The star-shaped leaves and corky bark ridges provide year-round textural interest, while the broad canopy offers substantial shade in suitable sites away from high foot traffic. Specific cultivars amplify these aesthetic qualities and address practical concerns. For instance, L. styraciflua 'Rotundiloba' features rounded leaf lobes instead of the typical pointed ones, along with deep red autumn coloration, making it a favored selection for specimen plantings in lawns or as street trees. This fruitless variety, first described in 1931 and propagated since 1968, reduces maintenance issues while maintaining the tree's visual appeal, though it may develop a narrow-angled structure prone to damage. Other cultivars, such as 'Moraine' with its upright form and reliable red fall color, further expand options for colder climates like . The resin of Liquidambar, known as storax, has a long history in as an expectorant and for treating ailments like coughs, ulcers, and skin issues. In , extracts from L. formosana fruits, including compounds like bertulonic acid, are employed to promote blood circulation and exhibit antihepatotoxic effects, such as reducing liver levels in experimental models. Native American communities, including the and , historically applied storax to wounds for its healing properties, as well as for controlling coughs and , often mixing the sap with other plants for effects. Storax also holds cultural significance, particularly in Asian traditions where it features in perfumery for its warm, balsamic scent derived from the resin's aromatic compounds, contributing to blends and fragrances. In regions of its native range, such as for L. orientalis, the resin has been valued since ancient times in practices and religious ceremonies for its fragrant properties. However, despite these benefits, the spiky, persistent fruits of fruit-bearing varieties pose limitations in urban settings, creating litter on hard surfaces like sidewalks and posing slipping hazards, which has led to preferences for seedless cultivars in maintained landscapes.

Timber and industrial applications

The wood of , the primary commercial species in the United States, is characterized by its moderate , with a Janka rating of 850 lbf, interlocked , fine and uniform texture, and an average dried density of 34 lbs/ft³ (545 kg/m³). These properties make it suitable for a range of applications, including production, manufacturing, furniture construction (particularly acoustic cabinets), interior trim, boxes, crates, and turned objects. Additionally, the is used for , crossties, , , and , contributing to its versatility in the hardwood industry. Resin extraction from Liquidambar species yields storax balsam, a fragrant sap obtained by damaging the bark, which has industrial applications in adhesives, lacquers, soaps, cosmetics, and perfumes. Historically, the hardened sap served as a natural , particularly among Native American communities and early settlers. In pharmaceutical contexts, storax from L. styraciflua has been incorporated into ointments and syrups, while L. orientalis provides a substitute used in and for similar purposes. L. styraciflua holds significant economic value in the lumber industry, where it supports production of , , and on a commercial scale across the . In , species such as L. orientalis contribute to exports of specialty woods and balsam, with storax valued in international markets for perfumery, pharmaceuticals, and manufacturing.

Cultivation practices

Liquidambar trees thrive in full sun exposure, requiring at least six hours of direct daily to promote vigorous growth and optimal fall coloration. They prefer moist, well-drained, acidic soils with a pH of 5.5 to 6.5, though they adapt to a range of types including clay and , and tolerate occasional wet conditions. Suitable for USDA hardiness zones 5 to 9, these trees are hardy in temperate climates but may experience winter injury in the colder extremes of zone 5. For mature specimens reaching 60 to 75 feet in height and 40 to 50 feet in spread, plant spacing of 50 feet (approximately 15 meters) is recommended to allow for full canopy development without competition. Propagation of Liquidambar is commonly achieved through , which require cold, moist at 41°F for 15 to 90 days to break and enhance rates, typically sown in after treatment. or stem cuttings, treated with rooting hormones, root successfully under high humidity and mist, providing a means to propagate selected cultivars, while onto is used for desirable traits like disease resistance. is the ideal time for initiating to align with the tree's active phase. Ongoing care involves moderate watering to maintain , particularly for young during the first two years, with established plants showing good once rooted deeply. should be done in late winter or early spring to shape the tree and remove dead or crossing branches, promoting a strong central leader and reducing disease entry points. Liquidambar exhibits tolerance to and compacted soils, making it suitable for landscapes, though it remains susceptible to diseases like Botryosphaeria, which cause trunk lesions and require prompt removal of infected tissue to manage spread. Key pests include bagworms, which construct protective leaf sacks and defoliate branches; small infestations can be controlled by hand-picking, while larger ones may necessitate applications. Although generally resistant to , some cultivars may show vulnerability under stress, with management focusing on and avoiding wounded planting sites. Post-2020 studies, including a 2025 analysis of urban trees in , highlight increased susceptibility to prolonged droughts in urban settings for , underscoring the need for supplemental during extreme weather. Drought and other stresses can exacerbate diseases and issues. Selected cultivars enhance cultivation options, such as and 'Oconee' for compact, shrub-like growth reaching only 10 to 15 feet tall, ideal for smaller landscapes, or 'Slender Silhouette' for narrow, columnar forms up to 50 feet high with minimal spread. Variegated varieties like 'Variegata' offer ornamental appeal with cream-edged leaves, while 'Rotundiloba' provides rounded lobes and fruitless habit to reduce litter. These selections are propagated vegetatively to preserve traits and improve adaptability in horticultural settings. Recent genomic research, including a 2024 chromosome-level assembly of L. styraciflua, facilitates breeding for improved traits such as disease resistance and climate adaptation. Hybrids with L. formosana demonstrate enhanced productivity for timber and bioenergy applications. The species has acclimatized successfully in regions like as of 2025.

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