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Lithospermum
Lithospermum
Lithospermum
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Lithospermum
Lithospermum purpurocaeruleum
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Boraginales
Family: Boraginaceae
Subfamily: Boraginoideae
Genus: Lithospermum
L. (1753)
Type species
Lithospermum officinale
Species[1]

84; see text

Synonyms[1]
  • Batschia J.F.Gmel. (1791)
  • Cyphorima Raf. (1819)
  • Lasiarrhenum I.M.Johnst. (1924)
  • Macromeria D.Don (1832)
  • Nomosa I.M.Johnst. (1954)
  • Onosmodium Michx. (1803)
  • Osmodium Raf. (1808)
  • Pentalophus DC. (1846)
  • Perittostema I.M.Johnst. (1954)
  • Psilolaemus I.M.Johnst. (1954)
  • Purshia Spreng. (1817), nom. illeg.

Lithospermum is a genus of plants belonging to the family Boraginaceae. The genus is distributed nearly worldwide, but most are native to the Americas and the center of diversity is in the southwestern United States and Mexico.[2] Species are known generally as gromwells or stoneseeds.

Taxonomy

[edit]

Plants of the World Online currently accepts 84 species.[3] Other sources accept about 50 [4] to 60 species in the genus.[5] A 2009 molecular study showed that the genus Onosmodium is included within Lithospermum.[2]

Species

[edit]

84 species are accepted.[3][4][6][7]

Formerly placed here

[edit]

Ecology

[edit]

Lithospermum leaves are eaten by the caterpillars of certain Lepidoptera, such as the moth Ethmia pusiella which has been recorded on L. officinale.

Uses

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The dried root of Lithospermum erythrorhizon is a Chinese herbal medicine with various antiviral and biological activities, including inhibition of human immunodeficiency virus type 1 (HIV-1).[9][10] Lithospermum erythrorhizon is native to Japan, where it has been traditionally used to make a purple dye. In southwestern North America, a species of this genus was used as a contraceptive by the Shoshone Native American tribe.[11]

Fossil record

[edit]

7 petrified nutlets and nutlet fragments of a Lithospermum species have been described from the Late Miocene Ash Hollow Formation, Ogallala Group, five km south of Martin in Bennett County, South Dakota. †Lithospermum dakotense sp. nov. shows similarities in size, shape, attachment and epidermal cell patterns to extant Lithospermum species. The fossil nutlets were preserved in various stages of maturity. The fossils closely resemble the nutlets of Lithospermum caroliniense and Lithospermum incisum.[12]

References

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

Lithospermum

View on Grokipedia
from Grokipedia
Lithospermum is a of flowering plants in the family , comprising 84 species of annual or perennial herbs and occasional shrubs distributed primarily in temperate and subtropical to tropical mountainous regions worldwide. Estimates of species number vary from 50 to 84 due to ongoing taxonomic revisions in . These plants are typically taprooted and hairy, with erect stems bearing cauline, sessile, entire leaves arranged alternately. Their inflorescences form panicle-like cymes or solitary flowers in upper axils, featuring deep-5-lobed calyces that enlarge in and 5-lobed corollas that are funnel-shaped or salverform, often or , with tube lengths exceeding the lobes. consist of 1–4 ovoid nutlets, 2.5–6 mm long, which are smooth to pitted or wrinkled, with basal attachment scars; the name derives from Greek words meaning "stone seed," alluding to these hard, stone-like nutlets. Native ranges span , , , , and parts of , with many species adapted to dry, rocky habitats. Common names for the genus include gromwell, puccoon, and stoneseed, reflecting their varied regional uses and appearances. Several species, such as L. erythrorhizon and L. officinale, are notable for their pharmacological potential, with roots containing pigments like shikonin used historically in traditional Asian, European, and Native American medicine for , , and wound-healing properties, as well as for producing red dyes in textiles and . North American species like L. canescens (hoary puccoon) have been employed by for dyes and as emetics or poultices. Taxonomically, Lithospermum exhibits significant diversity in floral morphology, , and chromosome numbers, with phylogenetic analyses revealing evolutionary patterns including and in some taxa.

Botanical Characteristics

Morphology

Lithospermum species are annual or herbs, subshrubs, or shrubs, typically reaching heights of 10-60 cm, though some can grow up to 90 cm, with erect or ascending stems that are usually hispid or strigose due to dense covering of stiff hairs. These stems often branch from the base and exhibit a rough texture from the hairs, which measure up to 1.5 mm in length with swollen bases. The leaves are simple, alternate, sessile or subsessile, and range from linear to lanceolate in shape, measuring 1-5 cm in length with entire or slightly revolute margins; they are frequently rough-textured owing to the presence of trichomes, including antrorse hairs up to 1.9 mm long. Inflorescences form as panicle-like cymes or solitary flowers in upper leaf axils, bearing flowers that vary in length from 5 mm to over 40 mm and exhibit colors ranging from white and yellow to blue in certain species, with five-lobed calyces and corollas that are funnel-shaped or salverform. Fruits consist of nutlets that are hard, smooth to pitted or wrinkled, 2.5–6 mm long, and often shiny and stone-like in appearance, contributing to common names such as "stoneseed" or "gromwell." Root systems are generally taprooted or fibrous, with some species developing tuberous roots; certain Asian species, such as , produce red pigments like shikonin in their roots.

Etymology

The genus name Lithospermum was coined by Carl Linnaeus in his 1753 work Species Plantarum, deriving from the Greek words lithos (λίθος, meaning "stone") and sperma (σπέρμα, meaning "seed"), in reference to the hard, stone-like nutlets produced by plants in this genus. This etymology highlights the 18th-century botanical emphasis on distinctive fruit characteristics as a key identifier for classification. Linnaeus designated Lithospermum officinale as the for the , a choice that anchored the name's application to species bearing these characteristic nutlets. Common names for Lithospermum reflect both European and Indigenous linguistic traditions tied to their physical traits and uses. In English, "gromwell" originates from the gromil (modern grémil), a compound of gré ("gray," referring to the seed color) and mil ("millet," alluding to the seed size and shape). "Stoneseed" directly echoes the , emphasizing the durable nutlets. Among North American Indigenous groups, "puccoon" derives from the Powhatan/Virginia Algonquian term poughkone, denoting plants used to produce red, purple, or yellow dyes from their roots.

Taxonomy and Phylogeny

Classification History

The genus Lithospermum was first formally described by Carl Linnaeus in 1753, in both Genera Plantarum and Species Plantarum, where it was placed within the family Boraginaceae and included six species distinguished by their smooth, white, lustrous, and erect nutlets, with L. officinale designated as the type species. During the 19th and early 20th centuries, taxonomic revisions expanded and refined the genus's circumscription, with key contributions from botanists such as Friedrich Wilhelm Lehmann (1818), who described species like L. strictum, and Augustin Pyramus de Candolle (1846), who broadened the genus to include diverse taxa now assigned to other genera like Buglossoides and Lithodora, dividing it into sections such as Eulithospermum, Rhytispermum, and Margarospermum based on nutlet and floral variations. By the 1920s, Ivan Murray Johnston initiated comprehensive revisions of Lithospermum and the tribe Lithospermeae (erected within Boraginaceae), describing 35 new species between 1924 and 1954 while segregating seven genera (e.g., Lasiarrhenum, Macromeria, Onosmodium) to address morphological heterogeneity, particularly in nutlet shape, corolla form, and indumentum; these efforts recognized approximately 50 species in the genus by the mid-20th century, amid ongoing debates over its placement in subfamilies like Boraginoideae, driven by interpretations of nutlet morphology and floral traits such as style insertion and pollen presentation. Significant mergers in the late 20th and early 21st centuries reshaped Lithospermum based on molecular phylogenetic evidence, notably the incorporation by James I. and Charles C. Davis of former genera such as Onosmodium (seven species, e.g., L. bejariense), Macromeria (11 species, e.g., L. exsertum), and Lasiarrhenum into Lithospermum, adding 18 species and affirming the genus's through analyses of DNA sequences and shared synapomorphies like erect, porcelain-like nutlets. This expanded the recognized species count to around 60, with further additions including three new species described by in 2018 (L. chihuahuanum, L. kelloggianum, L. tenerum), reflecting reductions in synonymy and new discoveries primarily in and the . Today, Lithospermum comprises 84 accepted species, centered in sensu stricto (s.s.) within the order , as per the IV classification (2016) and associated revisions emphasizing tribe Lithospermeae's cohesion via integrated morphological and genetic data.

Phylogenetic Relationships

Recent phylogenomic studies have resolved the evolutionary relationships of Lithospermum within the family, confirming its when including several previously recognized segregate genera such as Onosmodium, Macromeria, Lasiarrhenum, Perittostema, and Psilolaemus. This broad circumscription encompasses approximately 84 accepted species, addressing earlier uncertainties about potential based on limited sampling in pre-2021 analyses that suggested or non- for North American lineages. Key methodologies include target sequence capture of hundreds of nuclear loci (e.g., 298 regions) and genome skimming of complete genomes (78 protein-coding genes) combined with nuclear ribosomal DNA (nrDNA) ITS regions, yielding high support for major nodes with bootstrap values exceeding 95% and jackknife support of 93–100%. Within Lithospermum, species, such as L. officinale and L. afromontanum, form a basal grade, while species constitute a strongly supported monophyletic that diversified primarily in eastern and radiated northward and southward. Onosmodium species nest monophyletically within this clade, supporting its synonymization under Lithospermum, whereas former Macromeria taxa are scattered across the phylogeny but integrated into the expanded genus. The genus is closely related to other Lithospermeae members like Buglossoides, Glandora, and Aegonychon, with the tribe itself monophyletic based on plastome data. However, a 2025 study indicates that Lithospermum in the strict sense may be non-monophyletic, with the Central Asian genus Ulugbekia sister to or nested within it, though the combined receives full support (100% bootstrap). Divergence time estimates place the crown age of Lithospermum in the Mid-Miocene, approximately 14.5–11.3 million years ago, coinciding with climatic shifts that promoted diversification in arid and montane habitats. colonization occurred around 8.8–9 million years ago, with subsequent migrations to ~6.7–6.9 million years ago. Signals of hybridization are weak or absent in recent phylogenies, with topological incongruences between nuclear and datasets attributed primarily to incomplete lineage sorting rather than . These findings resolve longstanding debates on generic boundaries and provide a robust framework for understanding Lithospermum's within .

Distribution and Habitat

Geographic Range

The genus Lithospermum exhibits a , primarily concentrated in the , with approximately 84 accepted species worldwide according to recent assessments. The majority of species—estimated at around 70—are native to the , spanning , , and parts of , particularly in temperate and subtropical mountain regions. In contrast, the genus is sparsely represented in the tropics and entirely absent from . Centers of diversity are prominently located in and the , where over 40 species occur, including more than 30 endemics in alone; for instance, hosts 4 species within its . In , distributions extend to countries like , , , , and , though with lower species richness. The presence is limited to about 10 species in , such as L. officinale native to (from to ) and L. erythrorhizon in eastern ( and ), alongside minor occurrences in East and southern (e.g., , , ). Some species, like L. arvense, have been introduced outside their native range, appearing in and parts of such as southern . Most Lithospermum species have narrow geographic ranges, often endemic to specific ranges or localized habitats with extents under 1,000 km², contributing to their vulnerability; widespread exceptions include L. incisum across central and southern U.S. to . Recent analyses, including 2024 POWO data confirming 84 species, highlight ongoing taxonomic refinements. Climate modeling from 2025 predicts southward distributional shifts for Asian species like L. erythrorhizon in response to warming scenarios, potentially contracting northern ranges in while expanding in southern provinces.

Habitat Preferences

Lithospermum species generally inhabit dry to mesic environments, favoring well-drained soils such as sandy, gravelly, rocky, or substrates across a wide elevational range from to over 3,000 meters. These thrive in open or semi-open settings that provide ample , with many exhibiting suited to arid or semi-arid conditions. For instance, species like Lithospermum incisum occupy dry plains, hills, and rocky slopes in full sun, demonstrating resilience in low-moisture landscapes. In the New World, particularly North America, Lithospermum species are commonly found in prairies, open woodlands, savannas, glades, and desert-steppe habitats, often on limestone outcrops or in fire-prone ecosystems. Lithospermum canescens, for example, prefers dry prairies, rocky open woods, and upland savannas with sandy or loamy soils, occurring from low elevations to mid-montane zones. Similarly, Lithospermum multiflorum grows in sandy or gravelly open areas within ponderosa-oak and spruce-fir forests at elevations of 1,800 to 3,000 meters. These habitats reflect the genus's adaptation to disturbed or successional sites in temperate and subtropical regions. Old World representatives, such as , occupy grasslands, forest edges, hedgerows, and disturbed areas like meadows or rocky , predominantly on or basic soils. This is noted in European and temperate zones, including alpine meadows, where it tolerates rocky ground and quarries. favors grassy mountain slopes and hills in eastern , in well-drained loamy to clay soils under temperate climates. Overall, the shows a preference for neutral to alkaline (6.0–8.0) and exhibits some frost sensitivity in lower-elevation populations, though many are resilient in cooler, higher-altitude settings. Adaptations in arid-adapted New World species include narrower or succulent-like leaves to reduce water loss, alongside deep root systems that access subterranean moisture in drought-prone habitats like sagebrush steppes. In fire-maintained American prairies, species such as Lithospermum canescens benefit from periodic burns that clear and promote . These traits underscore the genus's in dynamic, resource-limited environments.

Species Diversity

Accepted Species

The genus Lithospermum currently includes 84 accepted , as recognized by (POWO) in 2024. These exhibit significant diversity patterns, with approximately 60 occurring in the Neotropics (primarily in montane habitats from to ), around 20 in the Nearctic region (spanning ), and about 4 in the Paleotropics (mainly in eastern ). Morphological variation is evident in flower colors, ranging from white and yellow to purple, and in nutlet shapes, which can be smooth, tuberculate, or angled, contributing to the genus's adaptability across habitats. Notable groups within Lithospermum include the puccoon-like species, characterized by bright yellow to orange flowers and often used historically for dyes, such as L. canescens in eastern . Another prominent group comprises Asian species with red roots valued for pigment production, exemplified by L. erythrorhizon. These groupings highlight the genus's biogeographic and utilitarian diversity. The following table enumerates selected major species, focusing on their native ranges and distinguishing traits:
SpeciesNative RangeDistinguishing Traits
L. officinale to and HimalayaPerennial herb with white tubular flowers; nutlets smooth and shiny; widely introduced elsewhere.
L. erythrorhizonEast to with purple flowers; red roots used for ; grows in temperate grasslands.
L. caroliniense and with orange-yellow flowers; hairy stems and leaves; known as hairy puccoon.
L. incisum and with yellow fringed corolla lobes; narrow leaves; adapted to dry prairies.
L. canescensEastern with golden-yellow flowers; hoary (hairy) foliage; roots yield .
Recent taxonomic updates, incorporating post-2021 molecular phylogenetic data from genome skimming and nuclear loci, have confirmed the of Lithospermum and stabilized the species count at around 80–84, up from earlier estimates of 50–70 by integrating clades and resolving synonymies.

Taxonomic Revisions

A pivotal taxonomic revision for Lithospermum occurred in 2009, when Weigend et al. utilized nuclear ribosomal ITS and chloroplast trnL-F sequence data, combined with morphological traits, to demonstrate that the North American genus Onosmodium (false gromwell) was nested within Lithospermum. This proposed merging Onosmodium into an expanded Lithospermum, transferring approximately 6 from Onosmodium and contributing to the inclusion of around 10 total when accounting for related segregates like elements of Macromeria and Nomosa. The merger rendered Lithospermum , addressing prior in the tribe Lithospermeae. Subsequent nomenclatural adjustments formalized these changes, with providing new combinations and synonyms to integrate Onosmodium species, such as Lithospermum bejariense for former Onosmodium bejariense.61[87:NCOLBA]2.0.CO;2) In , a 2018 revision by further refined species delimitation through detailed morphological study, reducing several taxa via synonymy; notable examples include Lithospermum albicans and Lithospermum calcicola as synonyms of L. incisum, and Lithospermum laevigatum under L. distichum, based on overlapping variation in nutlet verrucae and vegetative features. These reductions consolidated the Mexican flora, where Lithospermum now includes 43 , 32 endemic. Taxonomic revisions in Lithospermum have primarily employed integrated approaches, combining morphological characters—such as nutlet surface sculpturing (e.g., verrucae density and shape)—with molecular markers like ITS and trnL-F for delimitation. This has refined estimates by resolving cryptic synonymy and mergers, potentially adjusting global counts downward from earlier inflated figures. Controversies persist, particularly around the separation of Lithospermum caroliniense from morphologically similar segregates like former Onosmodium taxa, where subtle differences in corolla and nutlet traits fuel debate despite molecular support for inclusion. For Asian taxa, a 2025 phylogenomic study using genome skimming data reignited discussions on incorporating the monotypic Ulugbekia (U. tschimganica), resolving it as or nested within Lithospermum but noting unresolved positions due to limited sampling. Currently, Lithospermum is recognized with 84 accepted worldwide, encompassing approximately 20 recent synonyms from these revisions, as tracked in authoritative databases like (POWO) and Tropicos. These resources facilitate ongoing monitoring of nomenclatural shifts and support conservation efforts by clarifying species boundaries.

Ecology

Reproduction and Life Cycle

Lithospermum species exhibit a predominantly life form, with most taxa persisting for 3–10 or more years through a woody base or deep that enables resprouting after disturbance or seasonal . A few species, such as L. arvense, are annuals that complete their life cycle within one , relying on prolific production for persistence in disturbed habitats. This perennial strategy supports long-term population stability, as adult survivorship often exceeds 95% annually in stable environments like sand dunes, contrasting with higher mortality in juvenile stages. Flowering in Lithospermum typically occurs from spring to summer in temperate zones, spanning to July depending on and local , with hermaphroditic flowers featuring five stamens and a superior that develops into a of four nutlets. These tubular corollas, often white or yellow, attract pollinators and produce to facilitate visitation. is primarily entomophilous, mediated by bees and flies, though some species like L. caroliniense display with pin and thrum morphs that promote despite partial self-compatibility in cleistogamous flowers. is favored through spatial separation of anthers and stigmas, enhancing , while occurs in closed cleistogamous flowers as a reproductive assurance mechanism under low conditions. Seed dispersal in Lithospermum relies on or limited animal mediation, as the hard-coated nutlets lack specialized structures like hooks or wings but may be transported externally by mammals or internally by granivorous birds via endozoochory. The nutlets' durable pericarp contributes to seed longevity, remaining viable in banks for over five years, which buffers against unfavorable years. is triggered in spring following cold stratification to break , often requiring 60–90 days at 4–5°C, with optimal rates at 10–15°C after after-ripening during dry storage. Phenological patterns vary across habitats, with earlier flowering in arid regions—such as May onset for L. incisum in western North American prairies—compared to later peaks in mesic temperate areas, reflecting adaptations to and cues that align with availability and establishment windows. In semi-arid zones, reduced rainfall can shorten the flowering period, emphasizing the genus's responsiveness to environmental drivers in timing reproductive events.

Biotic Interactions

Lithospermum species engage in various biotic interactions that influence their survival and ecological roles. Herbivory primarily targets the vegetative parts of the plant, with leaves and flowers consumed by larvae of certain moths. For instance, the of Ethmia pusiella feeds on L. officinale, potentially reducing photosynthetic capacity and reproductive output in affected individuals. The hard, bony nutlets, often shiny and pitted, are generally avoided by herbivores due to their structural resilience, which limits and enhances dispersal potential. Pathogenic interactions include susceptibility to fungal rusts such as lithospermi, which infect leaves of species like L. canescens, causing and reduced vigor. Plants in the family, including Lithospermum, are susceptible to various viral s. Some species exhibit chemical defenses, with pyrrolizidine alkaloids in roots and tissues deterring further pathogen invasion or damage by exhibiting toxicity. Symbiotic relationships involve potential associations with arbuscular mycorrhizal fungi, which aid nutrient uptake, particularly , in nutrient-poor soils common to Lithospermum habitats; for example, L. canescens roots form such partnerships to support growth in sandy or disturbed environments. No evidence confirms nitrogen-fixing symbioses in the genus, distinguishing it from associates. In plant communities, Lithospermum acts as a in disturbed sites, colonizing open ground and stabilizing soils through rapid establishment, as seen in L. officinale on waste areas. It provides resources for native bees, supporting pollinators like those visiting L. caroliniense flowers during spring blooms. Certain species, such as L. purpureocoeruleum, exhibit through root exudates containing naphthoquinones, inhibiting nearby competitors in mixed microhabitats. Biotic threats include in rangelands, where heavy pressure reduces Lithospermum cover in bunchgrass communities, favoring less palatable species. While the genus has low overall invasive potential, weedy species like L. arvense (corn gromwell) can establish in agricultural fields, though they rarely dominate native ecosystems.

Human Uses

Medicinal Applications

Lithospermum species have been employed in across various cultures for their purported therapeutic properties, particularly and effects. In (TCM), the roots of (also known as Zi Cao) are used to treat conditions such as burns, sores, and due to their cooling and detoxifying actions. The plant's extracts are applied topically to promote and reduce , with historical records dating back to ancient texts for managing inflammatory skin disorders. In European folk medicine, the roots of Lithospermum officinale have been utilized as a and , often prepared as decoctions to alleviate urinary issues and gastrointestinal complaints. Indigenous North American groups, including the , have employed species like Lithospermum canescens (hoary puccoon) by brewing teas for external application to treat fevers accompanied by spasms. Additionally, Lithospermum ruderale (western stoneseed) was historically used by the and tribes as an oral contraceptive, with limited ethnographic records indicating its role in fertility regulation, though scientific validation remains sparse. The primary bioactive compounds in Lithospermum species are naphthoquinones, such as shikonin and alkannin, which exhibit properties by scavenging radicals and antimicrobial effects against like Staphylococcus aureus through membrane disruption and iron . These compounds contribute to the ' anti-inflammatory actions by modulating production and inhibiting pro-inflammatory pathways. Modern pharmacological research supports these traditional uses, particularly for L. erythrorhizon. A 2022 meta-analysis of 11 randomized controlled trials involving 1,024 participants demonstrated that TCM formulas featuring L. erythrorhizon as the principal herb significantly reduced (PASI) scores (mean difference = -2.00, 95% CI [-3.19, -0.80]) compared to controls, with enhanced efficacy in combination therapies and mechanisms involving IL-17A suppression and apoptosis promotion via shikonin. For antiviral activity, shikonin from L. erythrorhizon has shown inhibition of HIV-1 replication by blocking viral entry and downstream signaling, with up to 100% inhibition at non-cytotoxic concentrations in preclinical models. A 2024 review of TCM applications highlighted L. erythrorhizon's role in through Shiunko formulations, which accelerate re-epithelialization, formation, and synthesis while exerting antibacterial and effects. Safety concerns include potential from pyrrolizidine alkaloids present in some Lithospermum species, which can cause liver damage at high doses, as evidenced by elevated levels in animal studies and known risks in family plants. No Lithospermum-derived products are FDA-approved for medicinal use, limiting their application to traditional or investigational contexts.

Cultural and Industrial Uses

The roots of , known as purple gromwell or zi cao, have been a significant source of natural purple-red derived from shikonin pigments, historically used to color textiles and clothing in ancient and . This was particularly valued for staining fabrics of nobility and high-ranking officials, with records indicating its application in traditional practices dating back over a thousand years. In , it was employed by the Han people and ethnic minorities for ceremonial and everyday garments, symbolizing status and . Additionally, shikonin extracts from the roots have been incorporated into for their coloring properties, enhancing products like lipsticks and skin tints with natural pigmentation. Indigenous North American communities, including the and , utilized species such as Lithospermum ruderale (western stoneseed or puccoon) for cultural purposes, including as a plant to produce red and yellow pigments from the roots for basketry, , and body paint in traditional practices. In , Lithospermum officinale (European stoneseed) was associated with protective charms due to its hard seeds, believed to ward off ailments like stones, though such uses were more symbolic than widespread. Several Lithospermum species are cultivated as ornamentals in , particularly in rock gardens and xeriscapes for their low-growing, mat-forming habits and . For instance, Lithospermum diffusum thrives in well-drained, rocky under full sun, making it ideal for low-maintenance landscapes that conserve water. is typically achieved through seeds sown in early spring after stratification for better or by cuttings taken in summer, though success rates can be variable due to specific and light requirements. Industrially, Lithospermum extracts serve as natural pigments in eco-friendly dyes for textiles and , offering alternatives to synthetic options with benefits. However, large-scale remains limited owing to the plant's slow growth, low seed viability, and challenging germination, often necessitating or wild harvesting for commercial yields. Prior to the 20th century, L. erythrorhizon roots were actively traded along the from to and beyond, facilitating the exchange of purple dyes that influenced textile industries across regions during the Tang and later dynasties.

Evolutionary History

Fossil Record

The fossil record of Lithospermum is limited but provides evidence of the genus's presence since at least the , primarily through durable nutlets and endocarps rather than complete plants. In , the fossil record becomes more definitive during the . Lithospermum dakotense, a new species described from silicified nutlets in the Ash Hollow Formation (approximately 10 million years ago) of , closely resembles the nutlets of extant n species such as L. caroliniense and L. incisum in size (about 2 mm long), shape, epidermal cell patterns, and attachment structure. These nutlets, preserved in various stages of maturity, indicate a schizocarpic typical of the genus. Additional Tertiary records from include nutlets identified as Lithospermum from and deposits in , contributing to a broader pattern of diversification in central n grasslands during this period. No complete plants or vegetative remains of Lithospermum have been documented in the fossil record, likely due to the poor preservation of softer tissues compared to the robust nutlets. The Old World fossil record remains limited, with no confirmed Quaternary geological fossils reported, though archaeological contexts in Europe preserve L. officinale nutlets from Bronze Age sites.

Biogeographic Evolution

Lithospermum is estimated to have originated in during the , approximately 9.6 million years ago, following the broader Eocene radiation within the family. Phylogenetic analyses using nuclear and plastid genomes support this cradle, with subsequent long-distance dispersal events facilitating colonization of the around 7.8–9 million years ago, likely via the North Atlantic or rare transoceanic events rather than vicariance through the Bering . This single-origin hypothesis is bolstered by the of the genus, excluding minor incongruences from incomplete lineage sorting, which underscores a unified evolutionary trajectory from Eurasian ancestors to global diversification. Diversification within Lithospermum accelerated in the during the middle to , driven by across that promoted in xeric habitats, particularly in and the , where the genus exhibits its highest diversity. The uplift of mountain ranges, such as the Sierra Madre and the Rockies, further catalyzed clade by creating heterogeneous environments that isolated populations and fostered adaptive , with northern species expanding westward into montane niches. Pleistocene glaciations imposed range contractions, forcing relictual populations into refugia, followed by post-glacial recolonization that shaped current distributions without major lineage splits. Dispersal patterns in Lithospermum are characterized by rarity of long-distance events, attributable to the non-aerodynamic nutlets that primarily rely on barochory or limited endozoochory, limiting transcontinental jumps beyond the initial colonizations. Biogeographic reconstructions in , including Lithospermum, reveal no strong correlation between such dispersal successes and specific traits, emphasizing instead the role of climatic vicariance and opportunistic LDD in a complex, multi-modal framework. Independent dispersals to , as seen in L. afromontanum, highlight sporadic jumps, but overall patterns align with gradual radiation post-colonization. Projections under future scenarios indicate southwestward range shifts for certain species, such as L. erythrorhizon in , where the is expected to migrate approximately 800–860 km southwest by the 2050s under low- and high-emission pathways, accompanied by expansion into higher elevations amid warming temperatures. These modeled dynamics, integrating models with variables, suggest adaptive potential but underscore vulnerabilities to altered regimes in Asian refugia.

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