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Ginkgoopsida

Ginkgoopsida is a monotypic class of gymnosperm plants within the division Ginkgophyta, encompassing the single order Ginkgoales and represented in the modern era solely by the species Ginkgo biloba, commonly known as the maidenhair tree. This ancient lineage is characterized by deciduous, dioecious trees that can reach heights of up to 30 meters, featuring distinctive fan-shaped leaves with open dichotomous venation borne on both long and short shoots. The reproductive structures include microsporangia spirally arranged on branched axes and paired ovules on long peduncles that develop into seeds with a fleshy, malodorous outer layer containing butyric acid.^[1]^[2] The fossil record of Ginkgoopsida extends back to the Permian period, approximately 270 million years ago, with early forms appearing in the Triassic and achieving peak diversity during the Jurassic and Cretaceous periods, when numerous genera thrived across Laurasian continents including Asia, Europe, and North America. These extinct relatives exhibited varied leaf morphologies and reproductive features, but the group underwent significant decline following the Cretaceous-Paleogene extinction event, leaving G. biloba as the sole survivor by the Paleocene. Native originally to southwestern, southern, and eastern China, G. biloba is now widely cultivated globally in temperate and subtropical regions for its ornamental value, resilience to urban pollution and pests, and potential medicinal properties derived from its leaves and seeds.^[3]^[1]^[4] Notable biological traits of Ginkgoopsida include the presence of multiflagellated sperm cells, a primitive feature shared with cycads and ferns, which swim through pollen tubes to fertilize the egg, highlighting its evolutionary position as a "living fossil" bridging early seed plants and more derived gymnosperms. The class's unique combination of gymnospermous naked seeds and angiosperm-like leaf venation underscores its isolated phylogenetic status, with molecular and morphological studies placing it as sister to cycads (Cycadopsida), with this clade sister to conifers and other Pinopsida (including gnetophytes). Despite its relictual status, G. biloba demonstrates remarkable longevity, with individuals living over 1,000 years, and adaptability that has ensured its persistence amid dramatic climatic shifts.^[2]^[5]^[1]

Taxonomy and Classification

Definition and History

Ginkgoopsida is a class of gymnosperms proposed by Sergei V. Meyen in 1984 to encompass the order Ginkgoales, including both extant and extinct seed plant groups, while distinguishing it from other gymnosperm classes such as Cycadopsida and Pinopsida based on morphological and phylogenetic evidence from the fossil record.^[6] This classification emphasized unique features like the fan-shaped leaves and motile sperm in Ginkgoales, setting them apart within the broader gymnosperm lineage.^[6] Historically, ginkgophytes have been classified at the division level as Ginkgophyta, a monotypic group containing only the genus Ginkgo among living species, reflecting their isolated position among seed plants.^[7] Earlier systems, such as that of Adolf Engler and Karl Prantl in 1897, integrated Ginkgo into the order Ginkgoales within gymnosperms, recognizing its distinct family Ginkgoaceae while aligning it broadly with conifer-like groups. Meyen's proposal elevated it to class rank under the superclass Gymnospermae to better reflect evolutionary distinctions.^[6] In the taxonomic hierarchy, Ginkgoopsida serves as the class, with subclass Ginkgoidae, order Ginkgoales, family Ginkgoaceae, and genus Ginkgo as the sole surviving representative.^[6] The name Ginkgoopsida derives from the genus Ginkgo, introduced by Carl Linnaeus in 1771, combined with the suffix "-opsida," a standard botanical ending denoting a class-level grouping, rooted in Greek terms for appearance or form. This nomenclature underscores the class's focus on the characteristic morphology of ginkgophytes.^[6]

Phylogenetic Position

Ginkgoopsida, encompassing the order Ginkgoales, occupies a basal position among extant gymnosperm clades, recognized as one of three major classes (Cycadopsida, Ginkgoopsida, and Pinopsida, the latter including gnetophytes as subclass Gnetidae) in recent phylogenomic classifications.^[8] Phylogenetic analyses consistently place Ginkgoopsida as sister to Cycadopsida, diverging early in gymnosperm evolution and representing the earliest-branching clade within the group; this relationship is reaffirmed by recent phylogenomic studies as of 2024.^[9]^[2] This relationship is robustly supported by chloroplast phylogenomics, which resolve Ginkgo biloba as sister to cycads with high bootstrap values across amino acid and nucleotide datasets.^[9] Earlier molecular studies using 18S rRNA sequences also recovered a cycad-ginkgo clade as the basal gymnosperm lineage, though with moderate support (bootstrap 59%).^[10] Alternative placements, such as Ginkgoopsida sister to Pinopsida (conifers), appear in some older datasets but are contradicted by comprehensive nuclear and plastid phylogenomics.^[11] Cladistic analyses of morphological and molecular data affirm Ginkgoales as a distinct lineage within seed plants, with divergence from the cycad lineage estimated at approximately 325 million years ago during the early Permian.^[12] Mesozoic ginkgoaleans, analyzed through parsimony-based cladistics incorporating fossil taxa, exhibit derived traits such as biflabellate leaves and compound reproductive structures, supporting their isolation as a monotypic order among living forms.^[13] These analyses highlight Ginkgoopsida's persistence as a relict clade, with no close living relatives beyond this sister relationship to cycads.^[8] Debates persist regarding the monophyly of Ginkgoopsida when including extinct forms, particularly whether Permian taxa like Trichopitys heteromorpha represent stem-group members or core ginkgophytes.^[14] Recent cladistic parsimony analyses of Jurassic Ginkgoales favor a monophyletic grouping that incorporates Trichopitys, alongside genera such as Karkenia, Toretzia, Umaltolepis, Yimaia, and Ginkgo, based on shared ovulate structures and branching patterns.^[15] However, the basal position of Trichopitys—lacking fan-shaped leaves and featuring multi-ovulate axes—raises questions about whether broader Ginkgoopsida, as proposed to include such peltasperm-like ancestors, forms a strictly monophyletic class or a paraphyletic grade leading to crown Ginkgoales.^[16] Ginkgoopsida differs markedly from other gymnosperm classes in key anatomical features. Unlike Pinopsida, which possess resin canals in wood and leaves for defense and transport, Ginkgoopsida lacks these structures entirely, contributing to its distinct wood anatomy without oleoresin production.^[17] In contrast to Cycadopsida, characterized by pinnate or bipinnate compound leaves with circinate vernation, Ginkgoopsida features simple, fan-shaped leaves with dichotomous venation, a trait unique among gymnosperms and retained in the sole extant species Ginkgo biloba.^[18] These distinctions underscore Ginkgoopsida's isolated evolutionary trajectory within gymnosperms.^[2]

Morphology and Anatomy

Vegetative Features

Ginkgoopsida plants are characterized by a distinctive arborescent habit, typically manifesting as deciduous, dioecious trees that can attain heights of 20–35 meters in their extant representative, Ginkgo biloba.^[19] These trees exhibit monopodial growth with angular branching patterns, forming an often irregular crown with long, somewhat erratic branches that contribute to their resilience against wind and snow damage.^[20] A key feature is the presence of dimorphic shoots: long shoots bear spirally arranged leaves and support overall canopy expansion, while short shoots, up to about 7.5 cm in length, cluster leaves at their tips and are involved in vegetative reproduction through sylleptic branching.^[19]^[21] This dimorphism is evident in both living and fossil forms, enhancing adaptability in varied environments.^[19] The leaves of Ginkgoopsida are simple, flattened, and distinctly fan-shaped, with open dichotomous venation where veins fork repeatedly from the base without cross-connections, creating a parallel-like appearance.^[21] In G. biloba, the leaves are typically 5–10 cm wide, often bilobed due to a deep median notch, and borne alternately on long shoots or in dense clusters on short shoots; they are hypostomatous, with abundant stomata on the abaxial surface.^[21] Fossil representatives show morphological variations, such as the deeply dissected, multilobed leaves of Baiera, which resemble incised forms of modern ginkgo leaves but with fewer veins per segment (typically four or fewer), reflecting evolutionary diversity within the group.^[22] These leaves are petiolate, with slender, flexible petioles that facilitate fluttering in wind, and turn golden yellow before deciduous abscission in autumn.^[21] Wood anatomy in Ginkgoopsida features pycnoxylic secondary xylem, which is dense and composed primarily of tracheids, similar to that in conifers but evolved independently.^[19] Tracheid walls exhibit scalariform to opposite pitting on radial surfaces, particularly in early wood, aiding efficient water conduction without the need for vessels.^[23] Notably, the wood lacks resin ducts, distinguishing it from many other gymnosperms and contributing to its resistance to certain pathogens. This structure supports the tall habit while maintaining hydraulic efficiency. The bark of mature Ginkgoopsida trees is gray to brownish, becoming rough and deeply fissured with age, forming irregular ridges and plates that provide structural support and protection.^[21]^[24] Root systems are extensive and deep-rooted, often developing a taproot with lateral extensions that enhance stability and nutrient uptake; in native Chinese habitats, they are particularly adapted to well-drained, acidic loess soils, enabling survival in nutrient-poor, silty conditions.^[20]

Reproductive Structures

Ginkgoopsida, represented solely by the living species Ginkgo biloba, exhibit dioecious reproduction, with male and female reproductive structures occurring on separate individuals. This sexual dimorphism ensures cross-pollination between trees, a trait consistent across the class. Male trees produce microstrobili, or pollen cones, which are clustered on short shoots and release pollen grains containing multiflagellated sperm cells—a unique feature among extant seed plants, shared only with cycads. These sperm, numbering up to thousands per cell with numerous flagella, enable motility during fertilization.^[25]^[26] Female trees bear ovules in pairs on short, specialized stalks emerging from axils of spur shoots, without enclosing structures typical of angiosperms. Pollination is anemophilous, relying on wind to carry pollen from male to female trees, typically in spring. Upon landing on the ovule's pollination drop, pollen grains germinate, forming a pollen tube that grows through the nucellus toward the female gametophyte. Within this tube, the generative cell divides to produce the multiflagellated sperm, which swim short distances to the archegonia containing eggs—an archaic siphonogamous mechanism reminiscent of ferns and non-seed plants. Fertilization occurs in late summer or early autumn, often resulting in a single functional embryo per seed, though polyembryony can occur with multiple embryos developing from one fertilization event.^[27]^[28] Following fertilization, seeds develop naked on the stalks, lacking a true fruit but featuring a three-layered seed coat. The outermost layer, the sarcotesta, is a fleshy, orange-yellow tissue rich in butyric acid that emits a foul, rancid odor resembling vomit or rotting butter when mature or decaying, deterring some predators while potentially attracting others. Beneath it lies the hard sclerotesta for protection, and an inner spongy endotesta. Seed maturation occurs over approximately six months, from spring pollination to autumn dispersal, with dispersal primarily occurring in autumn via gravity as seeds drop from the tree, though animals such as birds and mammals may consume the sarcotesta and excrete the intact inner seed, aiding secondary dispersal. This dual mechanism balances local and wider distribution, though the odor limits animal involvement compared to other gymnosperms.^[25]^[29]^[30] In addition to sexual reproduction, G. biloba demonstrates asexual propagation through vegetative means, particularly via root suckers that emerge from adventitious buds on roots, forming clonal offspring genetically identical to the parent. This sprouting allows for natural colony expansion and is commonly exploited in cultivation for propagating selected cultivars, enhancing resilience in disturbed habitats. While rare, tissue culture techniques can also induce somatic embryogenesis from leaf or megagametophyte tissues, further supporting asexual cloning under controlled conditions.^[31]

Fossil Record

Paleozoic and Mesozoic Origins

The earliest fossil records of Ginkgoopsida date to the Late Paleozoic, specifically the Permian period approximately 290–270 million years ago (Ma), primarily in Laurasian regions such as France and Russia.^[32] Genera like Trichopitys, exemplified by T. heteromorpha from the Lower Permian of France, exhibit proto-ginkgoalean features including multiple-forked, non-laminar leaves and short, forking ovule-bearing appendages that resemble those in later ginkgophytes.^[3] Similarly, Psygmophyllum from Permian deposits in Europe and Asia displays bifurcating leaves with helically arranged structures, indicating early vegetative adaptations within the group.^[32] These sporadic finds suggest a modest initial presence in northern Pangaean floras, with limited taxonomic diversity during this era.^[32] Diversification accelerated in the Triassic period, marking a significant radiation of Ginkgoopsida beginning in the Middle Triassic around 245 Ma and peaking in the Late Triassic.^[33] Leaf genera such as Ginkgoites and Baiera emerged, characterized by wedge-shaped or fan-like laminae with dichotomous venation, alongside early reproductive structures showing morphological innovations like clustered ovules.^[32] Fossil evidence from Laurasian sites, including exceptionally preserved seedlings from the early Middle Triassic Voltzia Sandstone in France, reveals ontogenetic stages akin to modern Ginkgo, with bifurcating linear leaves and taproot development, underscoring the group's adaptation to terrestrial environments.^[33] This phase saw increased abundance in northern hemisphere floras, though reproductive fossils remained scarce compared to leaves.^[32] The Jurassic period represented the zenith of Ginkgoopsida diversity and distribution, with widespread occurrence across the Northern Hemisphere from approximately 200–145 Ma.^[3] Species assigned to Ginkgoites and Baiera, often distinguished by leaf segmentation patterns, dominate assemblages in Europe, North America, and Asia, reflecting a peak in morphogeneric variety with at least eight leaf types reported.^[32] Key fossil sites include the Yanliao Biota in northeastern China, dated to around 165 Ma in the Middle Jurassic Jiulongshan Formation, where well-preserved Ginkgo leaves (e.g., G. daohugouensis, G. parvifolia) alongside wood and ovulate organs provide insights into whole-plant reconstructions and paleoecological roles in warm-temperate forests.^[34] These records highlight the group's integral position in Mesozoic gymnosperm communities, with implications for broader seed plant phylogeny.^[32] Ginkgoopsida maintained high diversity into the Cretaceous period (145–66 Ma), particularly in the Early Cretaceous, when the group achieved its maximum generic diversity with over 15 genera recorded across Laurasian continents.^[3]^[16] Fossils of leaves, seeds, and pollen are abundant in formations such as the Dakota Sandstone in North America and the Yixian Formation in China, showing continued morphological variation including deeply dissected leaves in genera like Pseudocycas and Czebanoa.^[3] The Late Cretaceous witnessed the onset of decline, with reduced abundance in high-latitude floras due to increasing competition from angiosperms and climatic changes, setting the stage for post-extinction rarity following the Cretaceous-Paleogene boundary event.^[35]

Cenozoic Diversity and Decline

During the Paleogene period, Ginkgoopsida exhibited notable persistence across the northern hemisphere, particularly in Eocene floras where warm climates supported their distribution at high latitudes.^[35] Fossil leaves and fruits attributed to Ginkgo species, such as G. adiantoides, are common in deposits like the Green River Formation in the western United States, dating to approximately 50 million years ago, indicating diverse riparian and lacustrine habitats.^[3] These remains, preserved in oil shales and lake sediments, show morphological similarities to modern G. biloba, with fan-shaped leaves and ovule-bearing structures suggesting reproductive viability in subtropical to temperate environments.^[36] As global cooling progressed into the Oligocene, Ginkgoopsida shifted southward, maintaining presence in mid-latitude forests of North America, Europe, and Asia.^[35] The Miocene marked the onset of significant decline for Ginkgoopsida, driven by progressive climate cooling and increasing competition from diversifying angiosperms, which outcompeted them in ecological niches.^[35] Fossil records indicate a sharp reduction in abundance and geographic range, with Ginkgo disappearing from North American sites around 7–15 million years ago, as evidenced by the absence of leaves and reproductive organs in later Miocene sediments of the Pacific Northwest.^[3] In Eurasia, similar patterns emerged, with dwindling occurrences in floras from central Europe and eastern Asia, reflecting adaptation challenges to drier, more seasonal conditions.^[37] This period saw a transition from widespread dominance to relictual populations, underscoring the group's vulnerability to Neogene environmental shifts. By the Pliocene, Ginkgoopsida records became sparse but persisted in select regions, with the last widespread evidence in Europe and Asia around 5–2.5 million years ago.^[3] In Europe, G. adiantoides leaves were abundant in early Pliocene deposits but vanished by the late Pliocene, while in Asia, fossils from Japan and China document survival in temperate woodlands until the onset of Pleistocene glaciation.^[38] Near-total extinction occurred during the Quaternary, with no confirmed wild fossils beyond East Asia, leaving G. biloba as the sole survivor in isolated Chinese refugia, likely due to intensified cooling and habitat fragmentation.^[35] Complementary evidence from fossil wood and pollen reinforces this decline narrative. Ginkgoxylon, a form genus for ginkgophyte wood, preserves anatomical details like scalariform pitting and growth rings in Eocene (e.g., Clarno Formation) and Miocene (e.g., Vantage) petrifactions, highlighting structural continuity with living forms amid rarity.^[39] Ginkgo pollen, characterized by its boat-shaped morphology, appears consistently in Cenozoic sediments from the Paleogene through Pliocene, but frequencies drop sharply in Quaternary layers, signaling population crashes across continents.^[36] Such microfossils, often found in amber and lake cores, provide insights into dispersal and environmental tolerances during the final phases of diversification loss.^[35]

Living Representatives

Ginkgo biloba Description

Ginkgo biloba is the sole extant species in the class Ginkgoopsida, recognized as a deciduous gymnosperm tree typically reaching heights of 20–35 meters, though exceptional specimens in China exceed 50 meters.^[40] Its distinctive fan-shaped leaves, measuring 5–8 cm in width, emerge on long shoots and feature dichotomous venation, turning vibrant yellow in autumn before abscising.^[41] The tree exhibits dioecious reproduction, with male individuals producing catkin-like pollen cones approximately 2.5 cm long, while females bear pairs of ovules that develop into drupe-like seeds, each about 2.5 cm in diameter, enclosed in a fleshy, foul-smelling sarcotesta.^[42] Physiologically, G. biloba demonstrates remarkable resilience, being fully deciduous and cold-hardy down to -40°C, corresponding to USDA Zone 3 conditions.^[42] It shows strong tolerance to urban air pollution, heavy metals, and compacted soils, attributed to adaptations like reduced stomatal density and thickened mesophyll layers that mitigate pollutant uptake.^[41] The species is notably resistant to most insects and diseases, with few significant pests affecting mature trees, though its seeds contain ginkgotoxin (4'-O-methylpyridoxine), a neurotoxin that can induce convulsions if ingested in large quantities.^[40]^[41] Growth in G. biloba is characterized by an initial slow rate, accelerating to about 30 cm annually during early maturity, with individuals capable of living over 1,000 years—some documented specimens surpassing 2,500 years.^[41]^[40] The tree displays dimorphic branching, featuring long shoots for vegetative extension and short spur shoots that bear clustered leaves and reproductive structures, facilitating efficient resource allocation for cone and seed production.^[40] Genetically, G. biloba possesses a large diploid genome of approximately 10 Gb (2n=24 chromosomes), though polyploid variants including tetraploids have been identified in modern populations, contributing to high heterozygosity often maintained through clonal propagation via root sprouting.^[41] This genetic stability underscores its status as a living fossil, with morphology closely resembling Mesozoic ancestors.^[41]

Global Distribution and Cultivation

Ginkgo biloba, the only extant species in the class Ginkgoopsida, has a native range confined to southeastern China, where wild populations persist in scattered refugia across provinces including Guizhou, Anhui, Zhejiang, Chongqing, Hubei, and Hunan.^[41] These populations, often numbering fewer than 500 individuals per site, are small and genetically depauperate, with ongoing debate about whether they represent truly natural or anciently cultivated escapes; they are primarily found in mountainous regions at elevations of 300–1,250 meters, such as Mt. Dalou in Guizhou and Mt. Tianmu on the Anhui-Zhejiang border.^[41]^[43] In their natural habitat, G. biloba grows in warm-temperate mixed broadleaf forests dominated by deciduous and evergreen species, typically along flood-prone streamsides on slopes averaging 12–58 degrees.^[41] The species has been cultivated for over 2,000 years in China, initially for its edible seeds and later for ornamental and medicinal purposes, with early records from temple gardens in Japan and Korea dating back to the 11th century.^[44] European introduction occurred in the mid-18th century, with the first documented planting in England in 1754 by the Physic Garden at Chelsea, followed by widespread adoption across the continent for botanical collections.^[45] In North America, G. biloba arrived in 1784 via seeds sent to Philadelphia by William Hamilton, and it has since become a common street tree in temperate regions from the eastern U.S. to the Pacific Northwest.^[46] Today, it is globally distributed through cultivation, thriving in urban landscapes of Europe, North America, and even subtropical and tropical areas like parts of India and South Africa, where it tolerates a wide climatic range from USDA zones 3 to 8.^[47] G. biloba exhibits broad habitat adaptability, preferring well-drained, deep sandy or loamy soils in full sun, with a soil pH tolerance spanning 4.5–8.5, though neutral to slightly acidic conditions (pH 5.0–7.0) support optimal growth.^[41] Its resilience to environmental stressors, including air pollution, salt, drought, and soil compaction, makes it particularly suited to urban settings, where it often outperforms other trees in withstanding ozone and particulate matter.^[48] In cultivation, it succeeds on a variety of substrates from clay to poor urban fill, requiring minimal maintenance once established. Wild populations of G. biloba face significant threats from habitat fragmentation due to deforestation, agricultural expansion, and urbanization, which have reduced many native stands to isolated remnants vulnerable to genetic erosion.^[49] Overharvesting for seeds and bark in rural areas exacerbates these pressures, contributing to the decline of truly wild trees in southeastern China.^[49] Despite this, the species remains secure globally through extensive cultivation, with millions of planted trees in parks, avenues, and gardens ensuring its abundance outside native ranges.^[41]

Evolutionary Significance

Relation to Other Gymnosperms

Ginkgoopsida shares several fundamental reproductive traits with other gymnosperm classes, including the production of naked seeds not enclosed in an ovary and the presence of archegonia in female gametophytes, which are characteristic of non-flowering seed plants broadly. Like Cycadopsida (cycads), Ginkgoopsida exhibits flagellated, motile sperm cells delivered via zooidogamy, a primitive mechanism retained from fern-like ancestors, contrasting with the pollen tube-mediated siphonogamy in many conifers. Additionally, the wood structure in Ginkgoopsida, featuring pycnoxylic secondary xylem with distinct growth rings and lacking resin canals, shows similarities to that of extinct cordaites, an early gymnosperm group, and parallels aspects of conifer wood in its density and tracheid-based conduction.^[2] Despite these shared features, Ginkgoopsida possesses distinctive morphological traits that set it apart from other gymnosperms. Its leaves display open dichotomous venation, forming fan-shaped blades that fork repeatedly without cross-connections, unlike the linear, needle-like or scale-like leaves with parallel or reticulate venation typical of Pinopsida. Reproductive structures further diverge: Ginkgoopsida retains motile sperm without reliance on siphonogamy for delivery, a trait shared only with Cycadopsida among extant groups and absent in Gnetopsida, which employ non-flagellated sperm transported via elongated pollen tubes. The seeds of Ginkgoopsida are also unique in their bilateral symmetry and three-layered coat (sarcotesta, sclerotesta, endotesta), differing from the more uniform seed coats in conifers.^[50]^[51]^[2] In terms of evolutionary divergences, Ginkgoopsida lacks the compact, woody cones of Pinopsida, instead bearing loose, catkin-like strobili for pollen and ovules, and its broad, fan-shaped leaves contrast sharply with the pinnately compound fronds of Cycadopsida. Phylogenetically, Ginkgoopsida occupies a basal position within gymnosperms, often resolved as sister to Cycadopsida, with this clade branching before Pinopsida and Gnetopsida; molecular clock analyses support this topology, estimating the divergence of Ginkgoopsida from other extant gymnosperms around 300-350 million years ago in the Carboniferous. Some hypotheses propose Ginkgoopsida as sister to all other living gymnosperms, based on chloroplast and nuclear phylogenomics, though recent studies consistently favor its close alliance with cycads as the most deeply branching extant lineage.^[8]^[52]^[2]

Status as a Living Fossil

Ginkgo biloba, the sole extant species in the class Ginkgoopsida, exemplifies the concept of a "living fossil" through its remarkable morphological stasis, exhibiting minimal evolutionary change since the group's origins in the Permian period around 270 million years ago.^[53] This stasis is evident in the close similarity between modern G. biloba leaves and those of Jurassic fossils such as Ginkgo digitata, where fan-shaped laminae with dichotomous venation patterns remain virtually indistinguishable after over 170 million years.^[54] Such conservation extends to overall vegetative architecture, including the branching patterns and cuticle ultrastructure, which align closely with Mesozoic representatives.^[55] Key evidence for this status lies in the unchanged reproductive structures, particularly the presence of large, multiflagellated sperm cells that have persisted without significant modification for more than 200 million years, a primitive trait shared with ancient gymnosperm lineages. Leaf venation, characterized by open dichotomous branching without cross-connections, further underscores this continuity, as fossil imprints from the Triassic onward mirror the extant form in density and distribution.^[56] These features highlight a lineage that has maintained structural integrity amid profound environmental shifts, including mass extinctions.^[57] The survival of Ginkgoopsida can be attributed to several adaptive mechanisms, including the production of exceptionally long-lived individuals that can exceed 1,000 years in age, enabling persistence through climatic fluctuations.^[58] Clonal reproduction via basal lignotubers and embedded buds allows for vegetative sprouting and renewal after disturbance, circumventing seed-based vulnerabilities.^[59] Additionally, its broad physiological tolerance to temperature extremes and poor soils has facilitated endurance across diverse paleoenvironments, from subtropical Mesozoic forests to modern temperate zones.^[58] As a relict lineage with no close living relatives, Ginkgoopsida provides invaluable insights into paleobotanical reconstructions and evolutionary processes, serving as a direct link to gymnosperm diversity before the rise of angiosperms.^[5] This isolated phylogenetic position underscores its role in studying long-term stasis and resilience in plant evolution.^[58]

Human Uses and Conservation

Traditional and Modern Applications

Ginkgo biloba, the sole living species in Ginkgoopsida, has been utilized for millennia in traditional Asian practices, particularly in Chinese and Japanese contexts. The seeds, known as ginkgo nuts or ginnan, are consumed in Chinese and Japanese cuisine, often boiled, grilled, or incorporated into dishes like chawanmushi (savory egg custard) and rice preparations, valued for their nutty flavor and nutritional content.^[60] In traditional medicine, documented as early as 2800 BCE in Chinese texts, the leaves have been brewed into tea to address respiratory issues such as asthma and bronchitis, while boiled leaf preparations were applied topically for chilblains and other skin conditions.^[61]^[62] Seeds have similarly been employed in herbal remedies for cough, asthma, enuresis, and intestinal disorders.^[63] In modern applications, standardized leaf extracts like EGb 761, derived from Ginkgo biloba, are widely used for their purported benefits in cognitive enhancement and improving peripheral circulation, attributed to flavonoid and terpenoid components that promote vasodilation and antioxidant activity.^[64] Clinical trials, including randomized controlled studies, indicate that EGb 761 at doses of 240 mg/day can modestly improve cognitive function, neuropsychiatric symptoms, and activities of daily living in patients with mild to moderate dementia or mild cognitive impairment, though evidence does not support its role in preventing Alzheimer's disease or halting cognitive decline.^[65]^[66] Regulatory approval varies: in the European Union, EGb 761 is authorized as a herbal medicinal product for symptomatic treatment of cognitive disorders associated with aging and dementia, based on established use and clinical data, while the U.S. Food and Drug Administration has rejected health claims for ginkgo products, classifying them as unapproved new drugs without sufficient evidence for disease treatment.^[67]^[68] Beyond pharmacology, Ginkgo biloba serves ornamental purposes, prized for its fan-shaped leaves that turn vibrant golden-yellow in autumn and its resilience to urban pollutants, making it a common choice for street tree plantings in cities worldwide, including avenues in East Asia and parks in Europe and North America.^[41] The wood, though soft and lightweight, has been used traditionally for carving religious statues and crafting items like fans in Asian cultures.^[69] Fan-shaped leaves are also popular in bonsai cultivation and as a component in herbal supplements.^[70] Despite these applications, Ginkgo biloba seeds pose toxicity risks due to the presence of 4'-O-methylpyridoxine (MPN), an antivitamin B6 compound that can induce seizures, convulsions, and loss of consciousness upon overconsumption, particularly in children; cases of poisoning have been reported from ingesting 50-100 raw or undercooked nuts.^[63]^[71] Cooking reduces MPN levels, mitigating risks when seeds are prepared properly for culinary use.^[72]

Conservation Status

Ginkgo biloba, the sole living species in the class Ginkgoopsida, is classified as Endangered on the IUCN Red List due to its extremely restricted wild distribution and ongoing pressures on remnant populations.^[73] Wild populations are extremely rare and restricted to small, isolated stands in eastern and southwestern China, including sites in Zhejiang, Guizhou, and Hubei provinces, with individual populations typically comprising fewer than 100 trees.^[41] In contrast, cultivated populations are secure and widespread globally, with millions of trees planted in urban landscapes, parks, and plantations for ornamental and economic purposes.^[41] The primary threats to wild G. biloba include habitat destruction through deforestation and agricultural expansion in China, which has fragmented and reduced its native relic habitats to near extinction levels.^[59] Overharvesting for medicinal and ornamental uses has further depleted natural populations, while the proximity of extensive plantations poses risks of genetic hybridization that could erode the unique diversity of wild genotypes.^[59] Climate change exacerbates these issues by altering suitable habitats in montane regions, potentially shifting distributions and stressing remaining stands through increased temperatures and altered precipitation patterns.^[74] Conservation efforts focus on in situ protection within designated reserves, such as the Tianmushan Nature Reserve in Zhejiang Province, which safeguards one of the few semi-natural wild populations comprising 167 trees, many predating human cultivation.^[75] Ex situ strategies include seed banking, cryopreservation of germplasm, and propagation through tissue culture in botanic gardens worldwide to preserve genetic diversity.^[59] Despite these initiatives, the global outlook for wild G. biloba remains precarious, with low genetic diversity in surviving populations heightening vulnerability to environmental changes, though its abundance in cultivation ensures the species' persistence outside native ranges.^[41]

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Phylogeny of seed plants based on all three genomic compartments
Gnetales and conifers form a highly supported clade, and cycads are at the base of the gymnosperms, with Ginkgo branching next. Three gene-combined data ...
[12]
Divergence time estimation of Ginkgo in a gymnosperm-wide ...
Divergence between Ginkgo and cycads was estimated to ~ 325 mya, and the crown age of Ginkgo was 0.52 mya using the speciation birth-death.Missing: Ginkgoales cladistic
[13]
Phylogeny and evolutionary trends of Mesozoic ginkgoaleans
Meyen, 1984. S.V. Meyen. Basic features of gymnosperm systematics and phylogeny as evidenced by the fossil records. Bot. Rev., 50 (1984), pp. 1-111. View in ...
[14]
Permian ginkgophyte fossils from the Dolomites resemble extant O ...
Nov 3, 2010 · Trichopitys heteromorpha from the Lower Permian [13] is the oldest generally accepted ginkgophyte [[14], and other authors]. It consists of axes ...Missing: Ginkgoopsida monophyletic extinct
[15]
A new species of Karkenia (Karkeniaceae, Ginkgoales) from the ...
Aug 4, 2025 · The analysis favours a monphyletic group of Ginkgoales including Trichopitys, Karkenia, Toretzia, Umaltolepis, Yimaia and Ginkgo. The most ...<|separator|>
[16]
Review An overview of fossil Ginkgoales - ScienceDirect.com
Trichopitys, the oldest member so far known of this plant ... Contradictory opinions remain about the origin and phylogenetic relationships of Ginkgoales ...Missing: monophyly | Show results with:monophyly
[17]
Ginkgo | The Wood Database (Softwood)
Since ginkgo lacks resin canals, that can help separate it from both spruce and pine (which both have resin canals). Ginkgo's cell structure also tends to ...
[18]
2.6.2.1: Cycads and Ginkos - Biology LibreTexts
Jul 28, 2025 · 1 : Ginkgo leaves have a distinctive shape that has remained relatively unchanged from their fossil record: a fan-shaped leaf that is often ...
[19]
[PDF] Plant Guide
Ginkgo trees often have angular crown and long, somewhat erratic branches. They are usually deep rooted and resistant to wind and snow damage. Young trees are ...Missing: deciduous height dimorphic<|separator|>
[20]
Ginkgoales: More on Morphology
The fan shaped leaves of Ginkgo are flat and irregularly notched. They are often deeply grooved in the middle of the leaf, producing two distinct lobes.
[21]
None
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[22]
[PDF] analysis of circular bordered pit function ii. gymnosperm tracheids ...
In Ginkgo biloba and Taxodium distichum, the torus did not stain strongly ... Many of these species exhibited scalariform pitting, a geometry not ...
[23]
Ginkgo biloba - ResearchGate
The trees can live for hundreds of years. The bark is light to dark brown with rough grooves and reticulate fissures. The leaves are fan-shaped with bifurcated ...
[24]
Wake Up and Smell the Ginkgos - Arnold Arboretum
May 15, 2008 · Ginkgo Sexuality. Ginkgo biloba is a dioecious species, with separate male and female trees occurring at roughly a 1:1 ratio. Ginkgo shows a ...Missing: structures | Show results with:structures
[25]
Rethinking Ginkgo biloba L.: Medicinal uses and conservation - PMC
[2] The distinguishing feature of the ginkgo from conifers is its multiflagellated sperm cells, as in cycads, on vegetative anatomy, and partial molecular ...
[26]
[PDF] Ginkgo biloba - OpenSIUC
Mar 2, 2001 · Ginkgo is dioecious, meaning the male and female reproductive structures are produced on separate trees. These structures are restricted to the ...
[27]
The Diversity of the Pollen Tube Pathway in Plants - PubMed Central
Thus, pollen tube development of Ginkgo biloba shows an initial tubular multiaxial form (Figure 1) (Friedman, 1987), cycad male gametophytes typically show ...Missing: multiflagellated | Show results with:multiflagellated
[28]
Morphological Characteristics, Ultrastructure, and Chemical ...
Oct 13, 2023 · The mature Ginkgo biloba seed comprises five main parts: the sarcotesta, sclerotesta, endotesta, endosperm, and the seed embryo. Among these, ...
[29]
Ginkgo | Yale Nature Walk
Mar 3, 2016 · The pollen is dispersed via wind. Once the seeds fall off the tree in autumn and become fertilized, they can be dispersed by animals in order to ...
[30]
Gymnosperm Resprouting—A Review - PMC - PubMed Central
Nov 23, 2021 · Several reports of vegetative reproduction in fossil gymnosperms have been published, including epicormic resprouting and production of root ...
[31]
https://doi.org/10.1016/j.palwor.2009.01.001
[32]
https://doi.org/10.1016/j.palwor.2009.01.001
[33]
https://doi.org/10.1186/1471-2148-13-177
[34]
An evolutionary and cultural biography of ginkgo - Crane - 2019
Dec 4, 2018 · Ginkgo is also a living link to diverse seed plants that flourished over 100 million years ago. With recent discoveries of extinct fossil ginkgo ...
[35]
Tertiary Pollen-II The Oil Shales of the Eocene Green River Formation
Ginkgo or the Bennettitales. Four species or Arecaceae have been recorded ... Plant and insect fossils from the Green River Eocene of. Colorado. Proc ...
[36]
Ginkgophytes - ResearchGate
The ginkgophytes had their highest diversity and widest distribution in the Mesozoic, after which the group drastically declined during the Cenozoic. Today they ...Missing: Ginkgoopsida | Show results with:Ginkgoopsida
[37]
Ginkgo biloba's footprint of dynamic Pleistocene history dates back ...
Apr 27, 2018 · The genus Ginkgo first appeared in the middle Jurassic approximately 170 mya [1, 2]. As indicated by fossil evidence, a worldwide northern ...Missing: Ginkgoopsida characteristics
[38]
WOOD OF GINKGO - Botanical Society of America
In this paper, we discuss the anatomy of modern Ginkgo wood and of the Miocene Vantage and the Eocene. Clarno Ginkgo woods, and comment on other fossil woods ...
[39]
International Biological Flora: Ginkgo biloba - Lin - 2022
Feb 13, 2022 · Simulation of demographic history suggested that the lineage divergence of all living Ginkgo ... Based on DNA sequences, recent phylogenetic ...<|control11|><|separator|>
[40]
Ginkgo biloba - Oregon State Landscape Plants
Broadleaf deciduous tree, a gymnosperm, reaches 50+ ft (15+ m) tall, usually pyramidal, excurrent (dominant main leader), but variable.Missing: height angular dimorphic
[41]
Ginkgo - Maine Organic Farmers and Gardeners Association
... plant only male trees. However, the nuts are delicious and medicinal. Ginkgo was brought to England as an ornamental in 1754 and to North America around 1784.
[42]
Ginkgo biloba L. - Forest Service Research and Development - USDA
Ginkgo was introduced into North America in 1784 and has generally been successful on good sites in the moist temperate zone of the midwestern and eastern ...Missing: history | Show results with:history
[43]
[PDF] Ginkgo biloba Maidenhair Tree - Environmental Horticulture
Ginkgo tolerates most soil, including compacted, and alkaline, and grows slowly to 75 feet or more tall. The tree is easily transplanted and has a vivid ...
[44]
Ginkgo biloba, Ginkgo, Maidenhair Tree - UConn Plant Database
prefers deep sandy soils and moderate moisture; adaptable to pH and almost any other conditions; full sun; tolerant of pollution, salt air, and heat; prune in ...
[45]
Changes in Ginkgo biloba L.'s Habitat Due to Climate ... - MDPI
However, many Ginkgo populations are under threat from habitat destruction ... Given the increasing threats to wild Ginkgo biloba L. populations, exploring ...
[46]
[PDF] Male Gametophyte Development and Evolution in Extant ...
Two methods of sperm delivery occur in gymnosperms: zooidogamy, defined by pollen tubes with motile sperm as exhibited in cycads and Ginkgo, and siphonogamy, ...
[47]
Phylogeny and Divergence Times of Gymnosperms Inferred from ...
Sep 15, 2014 · Our study strongly supports cycads as the basal-most lineage of gymnosperms rather than sister to Ginkgoaceae, and a sister relationship between ...
[48]
[PDF] a true “living fossil” The Ginkgo - Oregon.gov
The trunk has light brown to brownish-grey bark that becomes deeply furrowed and highly ridged with age. Ginkgo leaves are fan-shaped, inspiring two more names ...Missing: fissured extensive loess
[49]
Lab IX - <EM>Ginkgo,</EM> <EM>Cordaites,</EM> Conifers (2)
Some ancient Ginkgo leaves closely resemble modern Ginkgo (VG 1:8). Others are highly dissected and resemble the typical ginkgo leaf but without tissue between ...
[50]
Comparative Studies of Leaf Cuticle Ultrastructure between Living ...
They found that the leaf cuticle ultrastructure of the fossil species is generally similar to that of the sole living ginkgolaean: Ginkgo biloba L. Both are ...Missing: venation | Show results with:venation<|control11|><|separator|>
[51]
Geometric and topological approaches to shape variation in Ginkgo ...
Both approaches also highlight that there are some fossils leaves that are very similar to modern Ginkgo leaves, and there are some fossil and modern leaves ...
[52]
Resequencing 545 ginkgo genomes across the world reveals the ...
Sep 13, 2019 · The model with the best scored supports the earliest divergence between the SWEST and EAST + SOUTH lineages being 515,780 years ago followed by ...
[53]
[PDF] Ecological conservatism in the "living fossil" Ginkgo
G biloba seeds above on left have been removed from their sarcotestas; those on the right have not. Paleocene Ginkgo seed on left (UWSP 14906) shows both ...
[54]
The modulation of immunological activities in human NK cells ... - NIH
Ginkgo seeds are used as a foodstuff in Japanese and Chinese dishes, confectionaries, and other foodstuffs, while ginkgo leaves are used as an herbal medicine ...<|separator|>
[55]
Synthetic and Natural Bioactive Molecules in Balancing the ...
Writings from 2800 B.C. describe its use as a memory enhancer and antioxidant powerhouse. Flavonoids and terpenoids in the leaf extract provide neuroprotection.
[56]
[PDF] Ginkgo Biloba Extract (GBE) - National Toxicology Program
The powdered leaf is inhaled for ear, nose, and throat disorders like bronchitis and chronic rhinitis. Locally applied boiled leaves are used for chilblains.
[57]
Review of Ginkgo biloba-induced toxicity, from experimental studies ...
The nut-like gametophytes inside the seeds have been used in traditional Chinese medicine for cough, asthma, enuresis, pyogenic skin infections, and intestinal ...
[58]
Ginkgo: Usefulness and Safety | NCCIH
Ginkgo has a long history of use in Chinese medicine for treating cough, diarrhea, and other conditions. Today, ginkgo leaf, usually as an extract, is the most ...
[59]
Efficacy and tolerability of Ginkgo biloba extract EGb 761® in dementia
The objective of this systematic review was to evaluate current evidence for the efficacy of Ginkgo biloba extract EGb 761® in dementia.Missing: enhancement | Show results with:enhancement
[60]
Expert consensus on the use of Ginkgo biloba extract, EGb 761 - NIH
In those with mild cognitive impairment (MCI), EGb 761® has also demonstrated significant symptomatic improvement versus placebo. World Federation of Societies ...
[61]
[PDF] Assessment report on Ginkgo biloba L., folium
Jan 28, 2014 · Chronic treatment with EGb 761 reduced the density of β-adrenoceptor (no subtype specified) binding to cerebral cortex and β-adrenergic- ...
[62]
[PDF] Scientific Memorandum: Ginkgo biloba extract (5/24/2012) - FDA
May 24, 2012 · Moreover, there is no food additive regulation in effect establishing safe conditions ofuse for Ginkgo biloba extract, and FDA is not aware ...
[63]
An evolutionary and cultural biography of ginkgo - Academia.edu
Ginkgo wood has been used to carve religious stat‐ Zhejiang Province (Gong, Chen, Dobes, Fu, & Koch, 2008; Gong, Qui, ues and old ginkgo trees are commonly ...
[64]
How to Grow & Care for Ginkgo Trees - Garden Design
Their pollution resistance and salt tolerance makes ginkgo trees a good choice for urban environments. Many cultivars have been bred for a smaller stature more ...Missing: industrial applications
[65]
4′-O-methylpyridoxine: Preparation from Ginkgo biloba Seeds and ...
It has been found that 4′-O-methylpyridoxine (MPN) is responsible for the poisoning caused by G. biloba seeds. The objective of this study was to explore and ...
[66]
In vivo toxicity assessment of 4'-O-methylpyridoxine from Ginkgo ...
Oct 15, 2021 · 4'-O-methylpyridoxine (MPN), a recognized antivitamin B 6 compound, is a potentially poisonous substance found in Ginkgo biloba seeds and leaves.
[67]
Ginkgo biloba: The tree that outlived the dinosaurs - Kew Gardens
May 5, 2020 · Evolutionary distinct and endangered Ginkgo biloba is rated as Endangered (EN) according to IUCN Red List criteria. The singular species has ...
[68]
Distribution pattern of large old Ginkgo biloba in China under ...
May 16, 2024 · Large old Ginkgo biloba trees (LOGTs), with profound ecological and cultural significance in China, face increasing threats from climate ...Abstract · INTRODUCTION · MATERIALS AND METHODS · DISCUSSION
[69]
The Ginkgos of Tian Mu Shan - Conservation Biology - Wiley
The Tian Mu Shan Ginkgo population is biologically significant by virtue of its long survival in a semi-natural state under conditions of intense interspecific ...