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Crepis

Crepis is a of flowering plants in the family , comprising approximately 200 of annual, biennial, and perennial herbs commonly known as hawk's-beards. These plants are typically 3–120 cm tall, with milky sap, taproots or rhizomes, and stems that are erect to decumbent, simple or branched, and often hairy. They feature basal rosettes and cauline leaves that are coarsely lobed or entire, and produce erect heads in cymiform, corymbiform, or paniculiform arrays, with 5–100+ yellow, orange, or sometimes white, pink, or reddish florets per head. The receptacles are epaleate, the involucres calyculate, and the cypselae are ribbed, curved, and either monomorphic or dimorphic, topped by pappi of barbellulate bristles. Native primarily to and , with about 12 species indigenous to (many polyploid with a base chromosome number of x = 11), Crepis species have been introduced nearly worldwide and can be found in diverse habitats from moist meadows to dry grasslands. Some species exhibit (asexual seed production), contributing to taxonomic complexity, while others are diploid or polyploid, reflecting evolutionary adaptations across their range. The genus is taxonomically challenging due to morphological variability and hybridization, as detailed in early cytogenetic studies. Notable for their dandelion-like appearance but distinguished by their pappus structure, Crepis species play roles in ecosystems as pollinator attractors and occasional weeds in agricultural settings. Certain taxa, such as Crepis intermedia, are used in restoration ecology for their adaptability to limestone soils and erosion control.

Taxonomy

Etymology

The genus name Crepis derives from the Ancient Greek word krepis (κρηπίς), which means "slipper," "sandal," or "boot." This term was used in ancient botanical texts, notably by Theophrastus in his Enquiry into Plants around the 4th century BCE, to refer to a plant, though the exact species is uncertain. Carl Linnaeus formally established the genus Crepis in his 1753 work Species Plantarum, where he described several species based on earlier observations. The name had appeared earlier in Linnaeus's 1737 Genera Plantarum, but the binomial nomenclature was solidified in 1753. The etymology likely alludes to the shape of the plant's achenes (cypselae) or the pappus, which may resemble a slipper or sandal base; alternative interpretations suggest a connection to the basal rosette of leaves or the foundational structure of the fruit.

Classification and phylogeny

Crepis is classified within the family , subfamily Cichorioideae, tribe (formerly known as Lactuceae), and subtribe Crepidinae. This placement reflects the genus's characteristic latex-bearing stems and composite inflorescences typical of the tribe. Historically, in the influential Genera Plantarum by Bentham and (1873), Crepis was included in the family Compositae (now ), tribe Cichorideae, emphasizing morphological features such as the ligulate corollas and pappus structure shared with other chicory-like genera. Babcock's seminal 1947 monograph further refined the infrageneric classification, dividing the genus into 27 sections based on a combination of morphological, cytological, and distributional data, providing a foundational framework for understanding its diversity. Modern phylogenetic analyses have significantly revised these views, revealing Crepis sensu lato as polyphyletic rather than monophyletic. A key study by Enke and Gemeinholzer (2008) utilized nuclear ribosomal ITS and plastidial matK sequence data to reconstruct relationships within , demonstrating that Crepis species form multiple distinct lineages more closely allied with other genera than to each other. For instance, the type species C. tectorum clusters with Mediterranean and Near Eastern taxa, forming a sister to another group of regional species, while a well-supported monophyletic encompasses Central Asian and North American members now placed in the genus Askellia (formerly Crepis sect. Ixeridopsis), isolated from the core of the genus. This suggests historical generic boundaries require adjustment, with some Crepis lineages showing affinities to genera like Tolpis (e.g., T. staticifolia nested within Crepis) and broader ties within tribe to genera like (subtribe Hieraciinae) and related taxa. Hybridization and have profoundly shaped the evolutionary history of Crepis, particularly evident in the North American agamic complex, where reproduction via predominates. Babcock and Stebbins (1938) first described this complex in their monograph on American Crepis species, highlighting how interspecific hybridization among diploids, followed by chromosome doubling, generated apomicts that form morphologically intermediate populations. Subsequent molecular studies confirm multiple origins of these polyploids, with limited between sexual and asexual lineages, underscoring the role of these processes in driving diversification and complicating phylogenetic resolution within the . These dynamics align with broader patterns in , where reticulate evolution via hybridization has obscured strict bifurcating phylogenies.

Diversity and species

The genus Crepis comprises approximately 200 of annuals and perennials distributed primarily across the , with significant presence in temperate and subtropical regions of Europe, , , and . The highest species diversity occurs in the circum-Mediterranean area, where over 100 are recorded, reflecting the region's role as a major center of origin and radiation for the genus. Notable examples include C. sancta, a self-compatible annual common across Mediterranean and known for its biform achenes; C. capillaris, a widespread Eurasian weed with variable numbers and a history of cytological study; and n taxa such as C. acuminata, a primarily apomictic with narrow, pinnately lobed leaves. In western , C. barbigera, C. intermedia, and C. occidentalis form part of an agamic complex involving polyploid apomicts derived from hybridization among sexual progenitors. Patterns of endemism are pronounced in insular settings, such as the , where species like C. tybakiensis—initially considered restricted to —extend to sites like ; and the , home to endemics including C. canariensis on . No exhaustive global species inventory exists, as taxonomic delimitations remain incomplete; regional treatments, such as the Flora of North America (documenting 24 species), and ongoing revisions highlight challenges from and hybridization that blur species boundaries.

Description

Morphology

Crepis species are annual, biennial, or perennial herbs, typically ranging from 3 to 120 in height, and produce a milky throughout the . They often form basal rosettes of leaves and exhibit a general habit superficially similar to dandelions, though distinguished by their erect stems bearing multiple flowering branches. The stems are usually erect to decumbent, numbering 1 to 20 or more per plant, and may be simple or branched, often striate, and ranging from glabrous to variously hairy, including hispid, setose, or stipitate-glandular pubescence. Leaves are alternate, with basal leaves often forming rosettes and cauline leaves reduced distally; basal leaves are petiolate with winged petioles, while blades are elliptic, ovate, lanceolate to linear, or spatulate to oblanceolate in shape, and margins vary from entire to dentate, serrate, toothed, or pinnately lobed, frequently lyrate or runcinate. The indumentum on leaves and stems can be glabrous, tomentulose, setose, hispid, glandular-setose, or tomentellous, contributing to variability in texture across . Roots in Crepis are predominantly taprooted, though some species are rhizomatous, with roots described as deep or shallow, woody or fibrous, and caudices often woody in perennials; annuals and biennials typically have shallower systems. This root architecture supports the erect, subscapose to caulescent , with stems arising from semi-woody rootstocks in some cases. Overall, the vegetative provides identification traits, emphasizing the combination of rosetted basal leaves, variable leaf dissection, and pubescence patterns.

Inflorescence and fruits

The inflorescence of Crepis species arises from branched scapes, typically bearing 5–50 capitula (flower heads) arranged in cyme-like, corymbiform, or paniculiform arrays, though solitary heads occur in some taxa. Each capitulum is erect and measures 1–2 cm in diameter, enclosed by a cylindric to campanulate involucre of phyllaries that is calyculate (with outer reduced phyllaries). The heads contain 5–100 ligulate () florets per capitulum, with corollas predominantly yellow but occasionally white, orange, pink, or reddish. All florets are morphologically bisexual, featuring a with sweeping hairs and anthers forming a around the style, but in apomictic species—which are common in the —they function as female, producing unreduced embryo sacs without and clonal seeds independent of . At the apex of each floret, a pappus of 80–150 white to tawny, barbellulate bristles in one or two series aids in wind dispersal of the mature fruits. The fruits are achene-like cypselae, monomorphic or dimorphic within a head, measuring 3–8 mm in length, and are subcylindric, , or curved with 10–20 that may be or roughened. Cypselae are typically yellow, brown, green, red, or black, with glabrous or hispidulous faces, and often taper to a (rostrum) at the ; the slipper-like shape of the cypsela base is thought to inspire the name from the Greek krepis, meaning "sandal." Unlike the dandelion (), which bears a single head per hollow scape, Crepis inflorescences feature multiple heads on solid, branched stems.

Distribution and habitat

Native range

The genus Crepis is native primarily to the , encompassing a broad distribution across (with a strong emphasis on the Mediterranean subregion), (extending from eastward to the and ), , and western . In , the genus is widespread from the and southward to the , , the , and the , as documented in comprehensive floristic surveys. North African occurrences are concentrated in the and , while Asian ranges include the , , the Altai-Tian Shan mountains, and extending into parts of and the . Native North American species are restricted to the and , representing a distinct evolutionary lineage within the genus. The center of diversity for Crepis lies in the and , where the highest and occur, with approximately 100 species confined to the . Specific hotspots include the , which harbor numerous narrow endemics such as Crepis tybakiensis, and the region, particularly and adjacent areas, supporting diverse assemblages adapted to montane and coastal habitats. These patterns reflect the genus's concentration in temperate to subtropical zones, with floristic data from regional surveys confirming over 40 taxa in alone, of which several are endemic. From a historical biogeographic perspective, Crepis likely originated in , specifically the Central Asian Altai-Tian Shan region, during the epoch, followed by westward dispersal and diversification. Major radiations post-dating the are evident in the Mediterranean, driven by climatic shifts and orogenic events that facilitated in fragmented habitats. This timeline aligns with and phylogenetic evidence indicating early establishment in Eurasian highlands before expansion into peripheral regions.

Introduced ranges and habitats

Several species of Crepis have been introduced beyond their native Eurasian ranges to regions including , , , and parts of , often via unintentional human-mediated dispersal such as contaminated crop seeds or wool shipments. For instance, C. capillaris (smooth hawksbeard) was first recorded in eastern and has since naturalized across much of the continent, while C. tectorum (narrowleaf hawksbeard) arrived before 1890 and occurs in at least 26 U.S. states, primarily in the northern half. In , both species are established, with C. capillaris noted as a weed since the late . C. capillaris is also widespread in , where it has been naturalized since 1867, and in , including since 1881. In introduced ranges, Crepis species typically occupy disturbed habitats such as roadsides, waste areas, grasslands, and agricultural fields, thriving in open, sunny conditions with poor to moderately fertile soils. C. capillaris favors lawns, turf, crops, and newly sown pastures in and , often forming dense stands in compacted or overgrazed sites. C. tectorum persists in similar settings across , including dry prairies and rangelands, and can overwinter as rosettes to exploit early-season resources. In Pacific regions, C. capillaris invades national parks like Haleakala in , adapting to volcanic soils and grasslands. Many introduced Crepis species exhibit invasive tendencies, competing with native plants for light, water, and nutrients, which can degrade biodiversity in grasslands and disturbed ecosystems. For example, C. tectorum invades lightly burned native vegetation in Alaska and the Pacific Northwest, reducing forage quality for wildlife and persisting in dispersed populations that outcompete forbs. C. capillaris is listed as invasive by the USDA in states like Alaska and appears on noxious weed inventories, where it forms monocultures that suppress native species in pastures and natural areas. As of 2025, global assessments highlight their expanding weed status, particularly in temperate regions, with ongoing monitoring needed for emerging threats in South American grasslands.

Ecology

Habitat preferences

Crepis species predominantly occupy open, sunny environments, including meadows, grasslands, rocky slopes, and disturbed sites, where they benefit from ample exposure and minimal from taller . These habitats often feature well-drained, nutrient-poor soils, with many tolerating a range of 6.0 to 7.5, as observed in common taxa like Crepis capillaris. Such preferences align with their adaptation to xerophytic conditions, enabling persistence in dry, rocky terrains across temperate and Mediterranean regions. The genus exhibits a broad altitudinal distribution, from in coastal and lowland areas to over 3000 meters in montane and zones, as exemplified by Crepis acuminata which ranges up to 3300 meters in western . In Mediterranean climates, species demonstrate notable through deep taproots and efficient water use, thriving in seasonal dry periods without supplemental moisture. This resilience extends to arid zones, where xerophytic traits like reduced leaf surface and durable seeds facilitate survival in low-rainfall ecosystems. Several Crepis taxa display ruderal characteristics, colonizing edges, roadsides, and abandoned fields, where disturbance creates favorable niches for establishment. For instance, Crepis sancta exploits such transient habitats in fragmented landscapes, underscoring the genus's opportunistic . Overall, these preferences reflect evolutionary adaptations to varied abiotic stresses, from to gradients, as detailed in foundational studies of the .

Biotic interactions

Crepis species engage in various biotic interactions that influence their survival, reproduction, and ecological roles within communities. Pollination is primarily facilitated by insects from multiple orders, including bees and flies, which visit the ligulate flowers for nectar and pollen. Studies on urban populations of Crepis sancta indicate that bumblebees (Bombus spp.) and solitary bees are key visitors, though their abundance can be reduced by air pollution, while syrphid flies (hoverflies) increase with greater green space availability. Butterflies are less commonly recorded as pollinators for Crepis, with observations often limited by habitat fragmentation, though general Asteraceae pollination includes occasional lepidopteran visits. Herbivory poses significant pressure on Crepis, with several taxa targeting leaves, stems, and reproductive structures. Larvae of the broad-barred white moth (Hecatera bicolorata) feed on foliage and buds of Crepis species, such as C. capillaris and C. vesicaria, contributing to defoliation in open habitats. The gall-forming fly Tephritis formosa specifically attacks flower heads (capitula) of hosts like Crepis virens and C. capillaris, inducing that disrupt seed production. and weevils are also prevalent herbivores; for instance, various aphid species colonize including Crepis, extracting sap and potentially vectoring viruses, while weevils like those in the genus Phyllobius chew on leaves and stems across European Crepis populations. These interactions can reduce plant fitness, particularly in dense stands. Mutualistic associations further shape Crepis ecology, notably through potential mycorrhizal partnerships that enhance uptake. Crepis biennis forms connections within common mycorrhizal networks (CMNs) with arbuscular mycorrhizal fungi, facilitating exchange with neighboring like Inula conyzae, though the presence of additional competitors can alter these dynamics and reduce benefits. Seeds of Crepis serve as a food resource for granivorous , integrating the genus into diets and aiding dispersal in some cases. As in disturbed habitats, Crepis contributes to stability by colonizing early successional sites, such as glacial moraines, where species like C. nana initiate vegetation cover that binds and prevents . This role supports subsequent trophic levels, providing for herbivores and for pollinators while fostering development for later-arriving plants.

Reproduction

Sexual reproduction

Sexual reproduction in Crepis primarily occurs through outcrossing facilitated by insect pollinators, such as bees, which are attracted to the yellow ligules of the florets. Many diploid species exhibit self-incompatibility or pseudo-self-incompatibility systems that prevent or limit self-fertilization, promoting genetic diversity by ensuring pollen from other individuals is required for successful seed set. The flower heads (capitula) of Crepis consist of numerous florets arranged in a structure that favors cross-pollination. All florets display protandry, where male-phase anthers mature before female-phase stigmas, reducing within a floret; additionally, outer florets often enter the female phase while inner florets are still in the male phase, further encouraging transfer between capitula. This temporal separation, combined with the open, composite , allows generalist to effectively vector across plants. Following successful and fertilization of ovules, Crepis produces viable achenes as , typically 50–100 per capitulum depending on and conditions. Central achenes are lightweight and equipped with a pappus—a feathery of bristles that aids dispersal—while peripheral achenes are heavier and lack a pappus, promoting establishment near the parent plant. set is higher under cross-pollination than selfing, reflecting the efficacy of mechanisms. Sexual reproduction dominates in diploid Crepis species, which are widespread across and , where they form the primary mode of propagation in native populations. This contrasts with apomictic reproduction prevalent in polyploid derivatives.

Apomixis

Apomixis in Crepis refers to asexual seed production through gametophytic mechanisms, primarily apospory, where diploid embryo sacs develop directly from somatic nucellar cells in the , bypassing and fertilization to yield clonal progeny genetically identical to the maternal parent. This process is particularly prevalent in polyploid lineages, where induces epigenetic deregulation of sexual genes, facilitating the shift to unreduced formation and enhancing reproductive assurance in or unstable genomic contexts. Apospory is the dominant mode in Crepis, often resulting in autonomous development alongside the , and is commonly observed in allopolyploids derived from interspecific hybridization. A notable example is the North American Crepis agamic complex, which encompasses sexual diploid species such as C. acuminata, C. occidentalis, and C. pleurocarpa, alongside apomictic polyploids like C. barbigera and C. intermedia, forming extensive swarms with over 100 distinct apomictic forms. These polyploids, typically triploid to octoploid, arise from multiple hybridization events among diploids, with stabilizing hybrid genotypes and enabling rapid diversification across western North American steppes and montane habitats. The complex originated from a single maternal lineage within the endemic North American section Psilochaenia, with phylogenetic evidence indicating polyploid formation and spread primarily within the region rather than multiple introductions from . Apomixis occurs predominantly among , though it is facultative in some taxa, allowing variable rates of within populations and contributing to despite clonal . In facultative apomicts, such as certain members of the agamic complex, embryo formation can mix aposporous and meiotic pathways, with apospory rates exceeding 78% in many ovules, which supports both clonal fidelity and occasional . This dual-mode capability underscores the role of in maintaining as an evolvable trait across the genus.

Reproductive interference and evolution

In the Crepis agamic complexes, reproductive interference primarily manifests as asymmetric interactions where from apomictic polyploids disrupts production in co-occurring sexual diploids. Apomictic individuals produce viable that competes with conspecific on sexual stigmas, often leading to the formation of inviable or low-fitness interploidy hybrids (e.g., triploids or pentaploids) that fail to recruit into subsequent generations. Additionally, this heterospecific can induce mentor effects, temporarily breaking down in sexuals and promoting self-fertilization, which further reduces outcrossed set and in mixed populations. Such interference is unidirectional, as apomictic ovules develop autonomously without requiring , rendering apomicts largely immune to from sexuals. These dynamics have profound evolutionary implications for Crepis, enabling to facilitate rapid colonization and spread in disturbed or marginal habitats where sexuals struggle with limitations or environmental . However, the clonal nature of apomictic reproduction constrains long-term , increasing vulnerability to pathogens and environmental shifts while relying on occasional for novelty. In agamic complexes like the North American Crepis group, apomixis drives by stabilizing polyploid hybrids derived from diploid sexual progenitors, resulting in morphologically diverse but genetically uniform lineages that blur species boundaries. Recent genomic analyses confirm that these complexes exhibit monophyletic origins with reduced heterozygosity in apomicts compared to sexuals, underscoring apomixis's role in fixing hybrid genotypes at the expense of variability. Over evolutionary timescales, reproductive interference may promote spatial of sexual diploids, which predominate in stable, low-disturbance environments where their higher confers a minority cytotype against pressures. This potentially allows for the restoration or maintenance of diploid sexual populations, countering apomict dominance and enabling that replenishes variation in complexes.

Uses

Culinary applications

Several species of Crepis are utilized in traditional Mediterranean cuisines, particularly as wild greens harvested for their leaves and young shoots, which are valued for their nutritional content and mild flavor when young. In , , the leaves of C. commutata and C. vesicaria are commonly gathered in spring and prepared raw in salads, boiled, steamed, or browned, often mixed with other greens to resemble in taste and texture. These preparations are integral to local dishes like boiled green mixes (horta) or savory pies (chortopita), with boiling employed to reduce inherent bitterness. Nutritionally, young shoots and leaves of C. vesicaria and C. commutata are rich in vitamins A and E (from and ), , and minerals such as calcium, magnesium, , and iron, contributing to their role in traditional diets for digestive and cardiovascular health. Harvesting focuses on tender growth to avoid the increased bitterness and tougher texture of older plants, which can make them less palatable without extended cooking. Ethnobotanical studies in Mediterranean regions highlight these species' importance in seasonal practices, where they provide essential micronutrients during periods of limited fresh produce. In , C. sancta is traditionally consumed as a wild green, known locally in regions like as "ciocapiat" or similar names evoking ("cicerbita" in some dialects), with young leaves boiled in mixtures or used in soups and salads. Similar to other Crepis species, older plants of C. sancta exhibit stronger bitterness, prompting warnings in ethnobotanical records to harvest early for optimal edibility. These uses reflect broader Mediterranean ethnobotanical traditions, where Crepis contributes to diverse preparations across and .

Ornamental and other uses

Several species of Crepis are valued in for their dandelion-like flowers and adaptability to dry conditions, making them suitable for rock gardens, borders, and meadows. Crepis rubra, an annual with striking pink florets, is particularly popular for its compact growth to 30 cm and low-maintenance nature, thriving in full sun and well-drained soils. It is often planted in cottage-style or informal gardens, where its solitary flower heads add color from summer onward. Crepis praemorsa, a bearing yellow flowers, is similarly employed in rock gardens for its once established, preferring gravelly, poor soils that mimic its native habitats. Beyond ornamentals, certain Crepis species serve as potential for , particularly in arid or disturbed areas where they grow as weeds. For instance, Crepis acuminata (tapertip hawksbeard) offers fair to very good value for sheep and , though its varies and it is often grazed opportunistically rather than as a primary . Species like Crepis tectorum are also consumed by in mixed pastures, contributing to without significant yield loss despite their weedy status. Medicinal applications of Crepis are limited and primarily rooted in traditions, with unverified efficacy. Crepis vesicaria has been used in ethnobotanical practices as an or to aid , alleviate , and treat , often prepared from its leaves. These uses persist in Mediterranean regions but lack robust clinical validation, emphasizing the need for caution in modern applications. Some Crepis species, such as C. intermedia, are used in for their adaptability to soils and .

Biochemistry

Secondary metabolites

Crepis species are known to produce a diverse array of lactones, predominantly guaianolides, which are characteristic secondary metabolites of the family. These compounds are commonly isolated from roots and leaves, contributing to the chemical profile of various taxa. For instance, crepin (C₁₄H₁₆O₄), a guaianolide-type , has been identified in the flower petals of Crepis taraxacifolia, where it occurs as a precursor that is activated upon . Other examples include 11β,13-dihydrozaluzanin C and its glycosides, reported from the roots of Crepis setosa, highlighting the prevalence of this skeletal class across the genus. Additionally, guaianolides such as those derived from zaluzanin C have been documented in the roots of Crepis rhoeadifolia, with epimeric forms exhibiting variations in stereochemistry at key positions. These are biosynthesized via the , involving as a key intermediate leading to the characteristic 5-7-5 ring system fused with a γ-lactone. Flavonoids in Crepis primarily belong to the luteolin-type, which are O-glycosylated derivatives concentrated in aerial parts. Luteolin-7-O-glucoside is a representative compound, detected in the leaves and flowering heads of multiple species, including Crepis tectorum and Crepis senecioides. This flavone glycoside features a luteolin aglycone with a glucose moiety at the 7-position, contributing to the flavonoid diversity observed in the genus. Other luteolin derivatives, such as luteolin 7-O-gentiobioside and luteolin 7-O-glucuronide, have also been identified in flowerheads, underscoring the chemosystematic significance of these compounds in Crepis . These arise from the phenylpropanoid pathway, with occurring post-flavonoid backbone formation. Phenolic compounds in Crepis include derivatives prevalent in aerial parts, with (5-O-caffeoylquinic acid) being widely distributed across species such as Crepis tectorum. This ester features a core acylated at the 5-position with , and it has been isolated from leaves and stems. Cichoric acid, a dimer of linked to , occurs notably in Crepis hokkaidoensis, where it predominates in the aerial biomass alongside caftaric acid. These phenolics are synthesized through the shikimate-phenylpropanoid route, yielding the characteristic hydroxycinnamic acid structures.

Nutritional and chemical properties

The leaves of Crepis species, particularly C. vesicaria subsp. taraxacifolia, offer notable , with dry weight compositions featuring 20-30% , primarily insoluble forms that support digestive and satiety in low-calorie diets. On a fresh weight basis, these leaves provide approximately 42 kcal per 100 g, alongside modest protein (1.04 g/100 g) and (0.69 g/100 g) contents, making them suitable for and nutrient-dense meal incorporation. Additionally, they contain vitamins, including β-carotene equivalents at 2-5 mg/100 g fresh weight and at 6-9 mg/100 g, which contribute to protection and . Chemically, Crepis leaves are rich in phenolic antioxidants, such as cichoric acid at levels up to 130 mg/g extract (equivalent to approximately 3% dry weight based on 23% extraction yield), which exhibit strong free radical scavenging activity in , ABTS, and FRAP assays, thereby reducing in biological systems. These compounds provide benefits, inhibiting production in macrophages with an of 0.428 mg/mL, without observed . lactones, common in the genus, serve as natural deterrents to herbivores by inducing in ruminants like sheep, yet overall remains low for human consumption. However, these lactones and the milky latex can trigger contact allergies or irritant in sensitive individuals, manifesting as upon direct exposure. Phytochemical analyses from 2020s studies highlight Crepis as a promising source of health-promoting metabolites, with cichoric acid concentrations comparable to or exceeding those in (0.5-1.5% dry weight), underscoring potential applications in functional foods for immune and antioxidant support.