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Capparis

Capparis is a genus of flowering plants in the family Capparaceae, comprising approximately 140–400 species (with 146 currently accepted) of shrubs, small trees, or climbers that are often spiny and distributed primarily in the tropical and subtropical regions of the Old and New Worlds. Members of the genus Capparis are characterized by simple, often leathery leaves, paired stipular spines (in many species), and flowers borne in axillary or terminal racemes or corymbs, featuring four sepals, four petals, numerous stamens, and an ovary elevated on a gynophore. The fruits are berries that are globular to ellipsoid, containing numerous seeds embedded in pulp. These plants thrive in diverse habitats, including arid and semi-arid environments, rocky cliffs, deserts, coastal forests, and tropical forests, often demonstrating strong adaptability with deep root systems. Under current classifications such as APG IV and POWO, the family includes about 15 genera and 500 , with Capparis being the largest . Traditionally, Capparaceae was circumscribed more broadly with about 45 genera and over 700 , but molecular data have shown it to be polyphyletic, with some elements now placed in or other families, though POWO maintains Capparaceae as distinct for the clade including Capparis. Taxonomically, Capparis belongs to the order and was first described by in 1753. distribution spans , (including the Mediterranean and Southwest ), the , and extends to temperate areas in southern and , with endemics in regions like and . Notable species include Capparis spinosa, the source of capers—its unopened flower buds pickled as a culinary in —and , valued in arid regions for its edible fruits and drought resistance. Many Capparis species hold ethnopharmacological significance, with traditional uses for treating , , and infections due to their content, including glucosinolates and ; modern studies support , , antimicrobial, and antidiabetic properties. Economically, they contribute to , medicine, and , particularly in tropical and subtropical .

Taxonomy

Etymology and Classification History

The genus name Capparis originates from the Latin capparis, borrowed from the Ancient Greek káppāris (κάππαρις), denoting the caper plant known for its edible buds. This nomenclature traces back to classical antiquity, with the term first documented by the Roman naturalist Pliny the Elder in his Naturalis Historia (circa 77 CE), where he describes the plant's properties and uses. The genus was formally established by in his in 1753, where he described several species and placed Capparis within the family , emphasizing its distinct floral and fruit characteristics. Early classifications maintained this placement in , with significant revisions in the 19th century by and in their Genera Plantarum (1862), who recognized subgenera and sections based on variations in fruit morphology and leaf structure, such as the sectional division into Capparis sect. Capparis and sect. Busbeckea. These morphological criteria guided 20th-century taxonomy, including a major revision by in 1965, which organized approximately 70 Old World species from to the Pacific into four sections, highlighting biogeographic patterns and resolving numerous synonyms. Further advancements in the late included regional monographs, such as those in the Med-Checklist project (Greuter et al., 1984) for Mediterranean taxa and Fici & Gianguzzi's 1997 study on diversity and conservation of wild and cultivated Capparis in , which discussed morphological variation using and field data. A pivotal shift occurred with the II classification in 2003, which subsumed into an expanded (s.l.) based on molecular phylogenetic evidence revealing close relationships, including shared biochemical traits like production. This reclassification underscored the of traditional and integrated Capparis into alongside relatives.

Phylogeny and Current Status

Recent phylogenetic studies have elucidated the complex evolutionary history of Capparis, employing molecular markers such as the ribosomal ITS , plastid matK , and extensive Angiosperms353 loci to reconstruct relationships across the . A 2025 focusing on taxa revealed polyphyly within sections like Monostichocalyx, alongside evidence of hybridization events inferred from whole-genome duplications and reticulate evolution, including multiple independent occurrences at the base of lineages and in Sect. Busbeckea. These analyses indicate at least five range expansions into , with rapid radiations and incomplete lineage sorting contributing to the genus's diversification. The study also describes five new species and two new , raising the total number of Capparis species in to 26. Under the APG IV classification system (2016), Capparis is situated in the core (s.s.) within the order , reflecting its distinction from expanded historical circumscriptions that included cleomoid and brassicoid lineages. However, molecular evidence continues to fuel debate over merging Capparaceae with , given their sister-group status and shared plastid traits, though APG IV maintains separation pending further resolution. The (POWO) database recognizes 146 accepted species in the genus as of its 2025 update, underscoring ongoing taxonomic refinements (noting pending incorporation of recent preprints). Infrageneric classification delineates four primary sections—Sect. Capparis, Sect. Sodada, Sect. Busbeckea, and Sect. Monostichocalyx—primarily based on reproductive traits: Sect. Capparis features indehiscent berry-like fruits, numerous stamens (30–150), and reniform brown seeds; Sect. Sodada is characterized by dehiscent siliquiform fruits opening along one suture, 6–15 stamens, and often winged or compressed seeds. Recent regional revisions have incorporated phylogenetic insights to propose new combinations; for instance, a 2022 study in () described the endemic C. kenaboiensis sp. nov. and two varieties of C. scortechinii, while a 2023 revision in PDR and introduced C. averyanovii sp. nov. and C. acutifolia subsp. thamphae subsp. nov., enhancing infrageneric clarity. Phylogenetic diversity hotspots in the Indo-Australian region, notably the Indochinese Peninsula as a key center and with its endemic radiations, guide conservation priorities by identifying evolutionarily distinct lineages for evaluations, such as vulnerable subspecies threatened by habitat loss.

Description

Morphology

Capparis species are typically woody shrubs, vines, or small trees with stems that can be erect, climbing, or sprawling, often reaching heights of 1–5 m or more in some taxa. The stems are frequently armed with paired stipular spines, which are straight or recurved and measure 1–10 mm in length, providing protection against herbivores; these spines may be absent or caducous on mature branches. Leaves are simple and alternate, usually distichous, with petioles 2–18 mm long; the blades are orbicular to ovate or elliptic, 0.5–10 cm long and 0.4–6 cm wide, often leathery or succulent in texture, with entire margins and a mucronate apex. The inflorescences are axillary, terminal, or ramiflorous, ranging from solitary flowers to racemose or paniculate clusters with 1–35 flowers. Flowers are bisexual and typically nocturnal, with pedicels 0.8–12 long; they feature four sepals in two whorls (outer often connate and saccate, inner free), measuring 3–25 mm, and four imbricate petals that are white to cream-colored (aging pink or ), oblong to elliptic, and 8–35 mm long. Stamens are numerous, varying from 6 to over 200 per flower, with filaments up to 5 long and anthers 1–3 mm; the gynophore is prominent, 1–8 long, supporting an inferior, 1-locular ovary with 2–10 carpels and numerous , topped by a sessile . Fruits are typically leathery berries borne on the gynophore and pedicel, ovoid to globose or , 0.8–5 cm long and 0.5–3 cm wide, with a smooth to warty or tuberculate pericarp that turns from green to red, orange, or purple at maturity; dehiscence is irregular or absent in most , though some exhibit capsule-like splitting. Each fruit contains 1–50 , which are reniform and 2–6 mm in diameter, embedded in a fleshy pulp or that is red, yellow, or white; the are dark brown to black, often tuberculate, and exhibit . Many Capparis display xeromorphic adaptations suited to arid or semi-arid environments, including thick cuticles on leaves and stems for retention, reduced size or succulence in taxa like C. spinosa, and dense indumentum of simple or stellate hairs that provides and minimizes . These features, combined with deep-rooted habits, enable survival in harsh conditions across tropical and subtropical regions.

Growth Habit

Capparis species exhibit a range of life forms adapted to diverse tropical and subtropical environments, predominantly as shrubs reaching 1-3 meters in height, though some tropical taxa develop into small trees up to 10 meters tall. For instance, Capparis tomentosa can form a small tree with an upright up to 15 cm in diameter or function as a spiny . Other species, such as those in tropical regions, include woody climbers or lianas that scramble or ascend host structures, enhancing canopy access in forested habitats. These plants display notable xerophytic adaptations, including deciduousness during dry seasons to conserve water, as seen in , where small, succulent leaves are caducous and persist for only about one month on new shoots. Coastal species demonstrate salt tolerance, thriving in sea-spray zones; Capparis spinosa var. inermis flourishes along shorelines, while Capparis cynophallophora exhibits high to saline conditions in dry coastal scrubs. systems further bolster , enabling access to subsurface water; Capparis spinosa possesses extensive roots that support growth during prolonged dry periods in Mediterranean ecosystems. Phenological patterns vary across the , with flowering typically occurring seasonally in response to environmental cues, often diurnal in many but nocturnal in others to attract specific pollinators. Fruiting follows flowering in a seasonal manner, aligning with favorable moisture availability for . species can achieve of several decades, contributing to their persistence in arid landscapes. Intra-generic variability includes spinose forms for defense against herbivores, contrasted with spineless variants like Capparis spinosa var. inermis, reflecting adaptations to differing ecological pressures.

Distribution and Ecology

Geographic Range

The genus Capparis exhibits a and subtropical distribution, spanning the , northern and eastern (with particular emphasis on ), western, central, and southeastern , , , and select regions of the , where species such as C. spinosa have been introduced; the genus is largely absent from temperate zones. This broad range reflects adaptations to arid and semi-arid environments, though populations are often restricted to coastal, rocky, or disturbed habitats. Centers of diversity for Capparis are concentrated in and , where over 50 species occur collectively, with Vietnam alone hosting 37 species and subspecies, many endemic. 's approximately 26 accepted species (including recent descriptions) further highlight this regional richness, contributing to disjunct distributions linked to Gondwanan vicariance and subsequent dispersal events originating from ancient African and Indian lineages. Seed dispersal in Capparis is primarily mediated by , which consume the fleshy fruits containing arillate seeds, facilitating long-distance spread across fragmented landscapes. Human activities have also influenced the genus's range, notably through the introduction of C. spinosa to Central and by Spanish colonizers in the , where it has since naturalized in suitable habitats. Habitat fragmentation poses significant threats to Capparis distributions, reducing and connectivity in remnant populations, as evidenced by studies on C. spinosa in arid regions. Recent assessments indicate that approximately 20% of are range-restricted, exacerbating vulnerability to land-use changes and impacts as of 2025.

Ecological Interactions

Capparis species exhibit diverse pollination strategies adapted to their often arid or semi-arid habitats, primarily relying on insect pollinators such as bees and moths attracted by nectar rewards and floral scents. In Capparis spinosa, for instance, diurnal bees and nocturnal hawkmoths serve as main pollinators, with intrafloral patterns of color and scent enhancing visitation by these insects. Crepuscular bees and hawkmoths are also key pollinators across multiple Capparis taxa in Mediterranean regions, where floral rewards like nectar are partitioned along ecological gradients to optimize interactions. Some tropical species, such as those in the Capparaceae family with similar floral traits, show evidence of bat pollination, though this is less common in the genus and typically occurs in low-density populations with large flowers. Many Capparis species demonstrate self-incompatibility, including late-acting mechanisms that prevent self-fertilization; for example, in Capparis jacobinae and Capparis retusa, self-pollinated flowers largely fail to set fruit despite pollen tube growth, promoting outcrossing and genetic diversity. Herbivory in Capparis is countered by a combination of physical and chemical defenses, with spines on stems and leaves deterring large mammalian browsers by restricting access and movement. These spines, prominent in species like Capparis spinosa and Capparis divaricata, function as a mechanical barrier, increasing time and energy costs for herbivores such as and bushbuck. Chemical defenses include glucosinolates, secondary metabolites that deter herbivores; in Capparis ovata and other species, the glucosinolate-myrosinase system produces toxic hydrolysis products upon tissue damage, contributing to an with specialized like Pierinae . Despite these defenses, fruits of Capparis species attract dispersers, facilitating seed spread; in Capparis sepiaria, birds such as sparrows consume the fleshy pulp, aiding dispersal while bypassing seed defenses. Symbiotic relationships in Capparis enhance nutrient acquisition in nutrient-poor soils, particularly through associations with in the . In Capparis spinosa, non-leguminous nitrogen-fixing microbes such as and Comamonas sp. colonize the zone, enabling atmospheric as demonstrated by acetylene reduction assays and growth in nitrogen-free media. These associative symbioses support plant establishment in arid ecosystems, where Capparis species act as pioneer plants, stabilizing soils through extensive systems and reducing erosion on slopes and disturbed sites. In semi-arid regions, such as those in and , Capparis shrubs contribute to by binding loose substrates and improving water retention, facilitating succession in degraded landscapes. Capparis species support biodiversity by serving as host plants for Lepidoptera and providing resources for avian communities. Various taxa, including Capparis spinosa, host caterpillars of pierid butterflies such as Pieris brassicae (large cabbage white) and Belenois java (caper white), offering larval food despite chemical defenses. Fruits and nectar attract birds, enhancing seed dispersal networks and supporting frugivorous species in dry forests. In Pacific islands, introduced Capparis populations exhibit invasive potential, outcompeting native flora in coastal and disturbed areas.

Human Uses

Culinary Applications

Capparis spinosa, commonly known as the caper bush, is primarily valued in culinary contexts for its unopened flower buds, harvested and preserved as capers, which serve as a tangy in . These buds are typically pickled in a of and or cured in sea salt to develop their characteristic briny, pungent flavor, enhancing dishes such as pasta sauces, fish preparations, and salads. The fruits of C. spinosa, referred to as caper berries, are larger and milder, often pickled whole and added to salads or served as appetizers for their subtle floral notes and texture. Preparation methods emphasize timing and preservation to preserve the buds' integrity; unopened buds are hand-picked before flowering, then either dry-salted for or submerged in , releasing mustard oils that contribute to their zesty profile. In Mediterranean traditions, capers are incorporated toward the end of cooking to maintain their shape and intensity, appearing in classics like puttanesca or seafood dishes. Historically, capers were used as a spice in and cuisines, documented in texts like the cookbook Apicius and by early botanists such as Dioscorides, who noted their role in flavoring and preservation techniques dating back over 2,000 years. Beyond C. spinosa, other species like find use in arid regions of and the , where its fruits are consumed raw, cooked as a in curries, or pickled for spicy flavoring in local dishes, while seeds add a bitter tang to traditional preparations. The fruits of C. decidua are particularly noted in for their role in pickles and vegetable sides, reflecting adaptation to desert environments. Nutritionally, Capparis species, especially C. spinosa, offer high levels of (providing up to 9% of daily needs per serving) and , alongside potent antioxidants such as , the richest natural source of which is found in capers, supporting their value as a flavorful yet healthful ingredient. Italy and Spain dominate global caper production and trade, with Spain reporting higher yields in its 2025 Andalusian harvest and and among the leading exporters of preserved capers, underscoring the ingredient's economic importance in Mediterranean . Modern culinary fusions occasionally incorporate capers into Asian-inspired , blending their brininess with regional spices for innovative condiments.

Medicinal and Other Uses

Capparis species, particularly C. spinosa, exhibit notable medicinal properties attributed to bioactive compounds such as glucosinolates, , and phenolic acids. Glucosinolates in C. spinosa contribute to effects by reducing paw and inflammatory markers in animal models, with hydromethanolic extracts demonstrating up to 69% inhibition at 400 mg/kg. activities are prominent, as evidenced by high phenolic content in leaf extracts yielding low values in DPPH assays, supporting potential in managing oxidative stress-related conditions like . Traditional uses in systems such as and include treatments for and skin ailments; for instance, root decoctions of C. spinosa and related species like C. sepiaria (Himsra) are employed for relief and dermatological issues, reflecting their role as agents in folk practices. Pharmacological studies further validate these applications through and limited clinical evidence. assays reveal activity, with extracts of C. spinosa fruits inhibiting bacterial biofilms by 70-79% against pathogens like Serratia marcescens at concentrations of 0.5-2 mg/mL. For antidiabetic potential, hydroalcoholic extracts reduce fasting blood glucose and HbA1c; a randomized double-blind placebo-controlled involving 54 patients (28 receiving the extract) showed an 11.2% decrease in fasting glucose (from 180 to 160 mg/dL) and 4.8% in HbA1c after 400 mg thrice daily for two months. These effects stem from enhanced insulin sensitivity and reduced , though human trials remain limited in scope and scale. Beyond medicinal applications, Capparis serves various non-culinary purposes. C. spinosa is cultivated as an for hedges due to its attractive foliage and flowers, enhancing landscapes in Mediterranean regions. Seed oils show biofuel potential, with extracts convertible to via , offering a viable alternative from arid-adapted species. Additionally, the extensive root systems aid in arid farming, stabilizing soil in drought-prone areas. Safety considerations include potential toxicity from unprocessed seeds, where glucosinolate produces isothiocyanates that may cause goitrogenic effects at high doses; however, extracts are generally safe with LD50 exceeding 4000 mg/kg and no observed at therapeutic levels.

Species Diversity

Accepted Species

The genus Capparis comprises 146 accepted species worldwide, with recognition based on a combination of morphological characteristics and molecular phylogenetic evidence that establishes distinct evolutionary lineages. These species reflect the genus's center of diversity in tropical and subtropical habitats from southern through and the Pacific islands. Among the accepted species, Capparis spinosa serves as the type species, native to the where it grows as a spiny in arid and coastal environments. Capparis decidua, distributed across the and parts of arid , is notable for its edible fruits, which are consumed fresh, pickled, or in traditional preparations and provide through high protein, fiber, and mineral content. In , Capparis mitchellii is a widespread inland species forming dense thickets, though regional populations face pressures from and are assessed as rare or vulnerable in certain subregions. Diversity within Capparis shows pronounced patterns of , including oceanic archipelagos in the Pacific and where isolation has driven . For instance, Capparis sandwichiana is endemic to the and listed as vulnerable owing to grazing, competition from non-natives, and coastal development. Identification of accepted species relies on basic dichotomous keys emphasizing traits such as leaf morphology (e.g., vs. ), inflorescence structure, presence or absence, and type across major sectional groupings like Sect. Capparis and Sect. Sodada.

Taxonomic Revisions and Formerly Included Taxa

Recent taxonomic revisions of Capparis have refined species boundaries through morphological, molecular, and ecological analyses, particularly in and . In , a recognized 11 , including the description of a new species, C. kenaboiensis, and two new varieties, C. scortechinii var. ruthiae and C. trinervia var. chungiana, all endemic to the region, based on and field examinations that addressed prior incomplete classifications. In , ongoing monographic work has described new species such as C. phatadke in , contributing to the resolution of taxonomic uncertainties in Indochinese habitats, though specific synonymy reductions remain part of broader regional checklists. A 2025 phylogenetic of Australian Capparis proposed five new (C. xylocarpa, C. megacarpa, C. loxophleba, C. lasiantha, C. splendidissima) and two new subspecies of C. spinosa (subsp. formicosa and subsp. insularis), along with elevating C. loranthifolia var. bancroftii to species rank as C. bancroftii, inferred from whole-genome duplication events, incomplete lineage sorting suggestive of hybridization, and multiple independent dispersals into ; these proposals, if accepted, would bring the Australian total to 26 . Several taxa formerly classified under Capparis have been transferred to other genera following phylogenetic re-evaluations, often due to differences in fruit morphology, inflorescence structure, and molecular markers. Transfers to Boscia include Capparis albitrunca becoming Boscia albitrunca, justified by capsule dehiscence patterns and woody habit distinctions, while some African and Asian species with elongate, indehiscent fruits were moved to Cadaba, such as elements of C. farinosa complex, emphasizing silique-like fruits over berried types typical of core Capparis. These post-APG exclusions (circa 2003 onward) narrowed the circumscription of Capparis s.s. to woody shrubs and lianas with indehiscent berries, excluding approximately 20-30 herbaceous taxa to Cleomaceae based on shared gynophore traits. Taxonomic challenges persist, including cryptic species in African lineages where morphological stasis masks , as seen in the C. spinosa group across and the Mediterranean, where 2024 analyses revealed fine-scale population differentiation potentially indicating undescribed taxa. In , a 2024-2025 revision process identified up to 20 previously undescribed entities through integrative phylogenomics, highlighting rapid radiations and polyphyletic sections like Monostichocalyx. These revisions have reduced the estimated number of accepted Capparis species from earlier broad counts of 250-400 to 146 globally as recognized by POWO, primarily through synonymy lumping in complexes like the C. spinosa group (now one species with two subspecies and four varieties, resolving numerous synonyms via uniform criteria across and ). This consolidation impacts , as narrower species concepts aid in prioritizing endemic taxa for protection amid habitat loss in tropical dry forests.

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