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Retama

Retama is a genus of flowering shrubs and small trees in the legume family Fabaceae, belonging to the tribe Genisteae, and comprising five accepted species native to arid and semi-arid regions of the Mediterranean Basin, North Africa, the Canary Islands, and the Middle East. These plants are characterized by their unarmed, straggling growth habit, slender and rush-like branches, reduced or deciduous leaves often replaced by photosynthetic cladodes, and clusters of white or pale yellow papilionaceous flowers that bloom in spring. Fruits are typically ovoid or cylindrical legumes containing one to several seeds. The most widespread species include Retama monosperma (bridal broom or white broom), distributed from southern and through to the eastern , and Retama raetam (white broom), which extends from the across the to the . Other species are Retama dasycarpa (endemic to ), Retama rhodorhizoides (restricted to the ), and Retama sphaerocarpa (native to , , northwest , and ). Retama species thrive in sandy, rocky, or coastal soils with low water availability, exhibiting high through deep taproots and efficient water-use strategies. They form symbiotic relationships with nitrogen-fixing soil bacteria, enhancing soil fertility and making them valuable for and in degraded arid lands. In addition to their ecological roles, Retama plants have significant ethnobotanical importance, particularly in traditional Mediterranean and North African medicine, where extracts from leaves, roots, and aerial parts are used to treat ailments such as , , , and infections. Pharmacological studies have identified bioactive compounds like , alkaloids, and , supporting activities including , , and effects. Ornamentally, species like R. monosperma are cultivated for their attractive blooms and are introduced in regions such as and , though some populations have become naturalized or potentially invasive.

Taxonomy

History and etymology

The genus Retama was established in 1838 by the American naturalist in his botanical work Sylva Telluriana, where he described it as comprising broom-like shrubs previously misplaced within other genera. This publication aimed to supplement earlier floras by introducing new genera and species overlooked or misclassified by contemporary botanists, with Retama positioned within the legume family (). The etymology of Retama traces to the Arabic term "ratam" or "retem," referring to desert broom shrubs, as transcribed by early explorers, and the related Hebrew "rotem," denoting similar arid-adapted plants mentioned in biblical texts. Rafinesque adopted this name to reflect the genus's characteristic morphology and , distinguishing it from broader broom taxa like . Early species descriptions predated the genus, with Swedish botanist Peter Forsskål naming the type species as Genista raetam in 1775 during his expedition to Arabia, published posthumously in Flora Aegyptiaco-Arabica; the specific epithet "raetam" directly derives from the vernacular. Throughout the , taxonomic revisions refined the classification, notably with the transfer of Genista raetam to Retama raetam by Barker Webb and Sabin Berthelot in 1842, as part of their des Îles Canaries, which emphasized morphological distinctions such as reduced leaves and elongated inflorescences. Historical synonyms for Retama include Boelia (proposed by Webb in 1853 in Otia Hispanica) and Lygos (introduced by in 1763 in Familles des Plantes), the latter rejected as a nomen rejiciendum under the International Code of Nomenclature for , fungi, and to preserve nomenclatural stability. These synonyms arose from early attempts to segregate broom-like based on geographic or floral traits, but subsequent revisions consolidated them under Retama.

Phylogenetic position

Retama is classified within the family , subfamily , tribe , and subtribe Genistinae. This placement reflects its position among the broom-like adapted to Mediterranean and arid environments. The genus comprises four species that share morphological traits such as erect shrubs with reduced leaves and papilionoid flowers, aligning it with other members. Phylogenetic analyses using nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) regions and chloroplast DNA (cpDNA) trnL-trnF intergenic spacers have demonstrated close relationships between Retama and genera such as , , and . Specifically, Retama clusters near subgenus Spartocarpus based on shared molecular signatures and morphological features like pod structure and arrangement. These studies confirm Retama as a monophyletic lineage within , distinct yet nested among these allied genera, supporting the tribe's overall excluding certain outliers. The evolutionary history of Retama is tied to the aridification of the during the , approximately 5–10 million years ago, coinciding with the that intensified dry conditions and promoted adaptations in leguminous shrubs. This period facilitated the diversification of taxa, including Retama, through selection for and nitrogen-fixing symbioses suited to nutrient-poor soils. A key synapomorphy for the tribe, including Retama, is the accumulation of quinolizidine alkaloids, which provide chemical defenses against herbivores and reinforce phylogenetic cohesion across genistoid legumes.

Description

Vegetative characteristics

Retama species are typically shrubs or small trees, growing to heights of 1 to 6 meters, with a graceful, spreading habit characterized by slender, flexible, and often drooping green branches that perform much of the due to the 's leafless periods. These branches are pubescent when young, with fine silky hairs, and lack thorns, allowing flexibility against and abrasion in arid environments. Leaves are greatly reduced or absent for most of the year, appearing as small, linear to lanceolate structures measuring 2–8 mm in length, simple, entire, and sericeous (silky-haired) on both surfaces to minimize water loss; they are quickly , shifting photosynthetic reliance to the green stems known as cladodes. This enhances survival in dry climates by reducing . The root system features a deep , often extending 10–20 meters or more in depth, supplemented by extensive lateral roots, enabling access to and uptake in nutrient-poor, sandy soils; these roots also form symbiotic nodules with nitrogen-fixing . Bark is thin and greenish-brown on younger stems, becoming grayish with longitudinal striations and peeling fissures as the matures, providing minimal while maintaining flexibility. The overall architecture, with its wiry, branching form and deep anchorage, suits Retama to stabilizing dunes against and .

Flowers, fruits, and

The flowers of Retama species are papilionaceous, typical of the family, featuring a composed of a large posterior standard petal, two lateral petals, and two anterior petals that enclose the reproductive organs. These flowers measure 5–16 mm in length, with white coloration predominant in species such as R. raetam and R. monosperma (often with purplish tips), though some exhibit pale yellow blooms. They occur in terminal or axillary racemes containing 1–15 flowers, supported by the slender, modified branches of the plant. The is 5-lobed and bilabiate, typically 2–4 mm long and green to purple in hue. The androecium consists of 10 stamens arranged in a diadelphous , with nine fused into a tube and one free. Flowering occurs from late winter through ( to in Mediterranean regions), with many species producing fragrant blooms to attract pollinators. The fruits of Retama are that vary slightly by but are generally small and indehiscent or tardily dehiscent, aiding in protection within arid environments. Pods are elliptical to ovoid or globoid, measuring 10–22 mm in length and 3–10 mm in width, with a to straw-colored pericarp that becomes leathery and wrinkled upon drying. Each pod typically contains 1–3 reniform or rounded , 3–8 mm in size, which are olive- to brown and smooth. Reproduction in Retama is primarily entomophilous, with nectar-producing flowers attracting bees such as Apis mellifera and other for . grains are and trizonocolporate, facilitating effective transfer by insect visitors. Seeds exhibit physical dormancy due to a hard, impermeable coat, which promotes longevity in soil seed banks where they remain viable for 5–20 years. rates are low without treatment (5–13%), but scarification—such as immersion in for 120 minutes—enhances success to 67–94%, particularly under optimal temperatures of 15–30°C and non-saline conditions. This mechanism ensures opportunistic establishment following environmental cues like rainfall in habitats.

Distribution and habitat

Geographic range

The genus Retama is native to the and adjacent regions, encompassing the , , , from to , the including and the , the , and the . This distribution reflects its adaptation to semi-arid and arid environments across Macaronesian and Mediterranean ecoregions, with the genus centered primarily in coastal and inland . Among the species, Retama raetam exhibits the broadest native range, extending from the through in , across the to the , and including southern in . In contrast, Retama monosperma is distributed in southwestern , particularly southern and including the , and northwestern in countries such as , , , , and . Retama dasycarpa is endemic to . Additionally, Retama rhodorhizoides is endemic to the . These patterns highlight disjunct populations shaped by historical climate fluctuations and barriers like the , which have isolated lineages in semi-arid zones. Introductions of Retama species outside their native range are limited and often purposeful. In , R. raetam was introduced as an ornamental in around 1841 and has naturalized in Mediterranean-climate regions, while R. monosperma is also established there. In , R. monosperma has been planted for in coastal and grasslands but has become naturalized and invasive in areas like County. Such plantings leverage the genus's utility in stabilizing sandy soils, though they pose risks of spread in similar dry environments.

Preferred environments

Retama species thrive in a range of arid to semi-arid climates spanning Mediterranean to conditions, where annual rainfall typically ranges from 100 to 500 mm, with some taxa enduring even lower in hyper-arid zones (25-100 mm annually). These exhibit broad , surviving extremes from -5°C to 45°C, with adaptations enabling persistence in regions featuring hot summers and mild to cool winters. The genus prefers well-drained soils such as sandy, gravelly, saline, or types, often with low and nutrient content, and a range of 6 to 8. Retama demonstrates notable tolerance to up to 10 dS/m electrical , as well as heavy metal contamination including and , allowing establishment in otherwise inhospitable substrates. These shrubs commonly associate with shrublands, coastal dunes, and systems, occurring at elevations from to 1500 m, where they contribute to stabilizing sandy or eroded landscapes. In terms of water relations, Retama species are drought-deciduous in extreme , rapidly shedding leaves to conserve resources while maintaining high water-use efficiency through efficient stomatal regulation and deep root systems. Retama has shown success in human-modified habitats, particularly for revegetation of degraded lands, mine tailings, and coastal stabilization projects, leveraging its to pioneer nutrient-poor and disturbed sites.

Ecology

Symbiotic relationships

Retama species, as members of the family, form symbiotic relationships with nitrogen-fixing , primarily alpha-proteobacteria such as Ensifer aridi, which induce the formation of root nodules. These nodules enable biological , allowing the plants to convert atmospheric N₂ into usable forms, with studies on Retama monosperma showing effective nodulation in arid and contaminated soils. In Retama raetam, this contributes 46–86% of the plant's , supporting in nutrient-poor environments and enhancing . Additionally, Retama associates with arbuscular mycorrhizal fungi (AMF), which colonize the to improve uptake from impoverished arid soils. Molecular analyses of Retama raetam in southern reveal diverse AMF communities, including genera like Glomus and Acaulospora, that enhance nutrient acquisition and plant resilience in low-fertility habitats. These associations are crucial for Retama's establishment in semi-arid ecosystems, where soil is often limiting. Pollination in Retama is primarily entomophilous, facilitated by insects such as bees that access floral rewards of nectar and pollen from the papilionaceous flowers. While specific pollinators vary by region, generalist bees and other insects ensure cross-pollination, aiding reproduction in sparse desert flora. Flower morphology, including the keel structure, supports effective pollen transfer during insect visits. Retama experiences herbivory from browsing animals like and camels in arid rangelands, where its foliage serves as , but quinolizidine alkaloids in the act as chemical defenses to limit excessive consumption and prevent . These shrubs provide during dry seasons, though is deterred by the alkaloids' bitter and potential health risks. Complementing this, Retama exhibits allelopathic effects through root exudates containing specialized metabolites, such as quinolizidine alkaloids and , which inhibit the of nearby competitor and parasitic weeds in arid zones. This interaction helps Retama maintain dominance in resource-scarce environments.

Environmental adaptations and threats

Retama species have evolved physiological mechanisms to cope with the harsh abiotic conditions of their arid and semi-arid habitats. Osmotic adjustment through accumulation is prominent, particularly in Retama raetam, where proline levels rise dramatically—up to 24-fold in drought-stressed subspecies—to maintain and protect cellular structures. This response is complemented by stomatal regulation, which minimizes water loss via partial or full closure during water deficits, thereby conserving resources while sustaining minimal . In saline environments, certain species exhibit tolerance via osmotic regulation, allowing survival on coastal or inland salty soils. These adaptations collectively enable Retama to thrive in low-water, high-salinity settings where other plants falter. Resilience to biotic and abiotic disturbances further bolsters Retama's persistence. Fire tolerance is achieved primarily through resprouting from lignotubers or basal buds following moderate-intensity burns, facilitating rapid post-fire and canopy regeneration in fire-prone Mediterranean ecosystems. Similarly, like Retama sphaerocarpa demonstrate from ; after prolonged heavy pressure, reduced allows regrowth and restoration of cover, highlighting their capacity to rebound in disturbed rangelands. Symbiotic with enhances this by improving nutrient availability in impoverished soils, supporting overall vigor under . Despite these adaptations, Retama populations confront significant anthropogenic and environmental threats. Habitat fragmentation from and disrupts connectivity in Mediterranean and North African landscapes, isolating remnants and hindering . Climate change intensifies aridity through prolonged droughts and rising temperatures, straining adaptive limits and potentially shifting suitable habitats beyond current ranges. Competition from , such as in coastal dunes where exotics outcompete Retama during dry periods, exacerbates local declines. Overharvesting for fuelwood in arid regions further depletes stands, particularly in areas with limited alternatives. Conservation efforts reflect Retama's varied risk profiles, with most species rated Least Concern by the IUCN due to their wide distributions, though localized threats prompt targeted actions. For instance, Retama rhodorhizoides faces localized threats from habitat loss on the Canary Islands. Many populations benefit from protection within nature reserves, such as those in Spain and Tunisia, where grazing controls and restoration initiatives safeguard key sites. Retama's utility in ecological restoration underscores its potential amid threats. Notably, Retama sphaerocarpa excels in phytoremediation of heavy metal-contaminated soils from mining spills, stabilizing substrates, accumulating trace elements like zinc and lead in roots, and reducing bioavailability to prevent leaching.

Species

Accepted species

The genus Retama comprises five accepted , as recognized by the database, all of which are shrubs adapted to arid and semi-arid environments in the and adjacent regions. These species are distinguished primarily by flower color (ranging from white to yellow), pod morphology (e.g., cylindrical versus spherical), and degree of leaf retention (most are deciduous or nearly leafless, with persistent needle-like leaves in some). Recent molecular studies, including inter-simple repeat (ISSR) fingerprinting and phylogenetic analyses of housekeeping genes, have confirmed the taxonomic boundaries among these , particularly resolving distinctions within North African populations. Retama raetam (Forssk.) Webb & Berthel., the , is a widespread reaching 2–6 m in height, with slender, drooping branches and small white flowers borne in axillary racemes. It features cylindrical pods and is nearly leafless for much of the year, with brief retention of small, linear leaves. Native from the Mediterranean coast through the to the and (including northern , , , and ), it thrives in sandy and rocky desert habitats. Retama monosperma (L.) Boiss. is a larger , typically 3–6 m tall (up to 10 m in optimal conditions), characterized by arching, rush-like stems, white pea-like flowers with purple sepals, and single-seeded pods that give it its name. It is nearly leafless, with occasional small linear leaves. Distributed across the (southern and southwestern ), , and extending to and the eastern , it prefers coastal dunes and dry shrublands. Retama sphaerocarpa (L.) Boiss., notable for its yellow flowers in short racemes and distinctive spherical pods (1–2 cm diameter), grows as an upright to 1–3 m (occasionally 4 m). It has minimal leaf retention, with small, early- linear leaves. Endemic to southwestern Iberia ( and central to southern ) and northwestern (from to ), it inhabits sandy soils and coastal areas. Retama rhodorhizoides (Webb & Berthel.) Webb & Berthel. is a semi-evergreen 2–4 m tall, with persistent needle-like leaves and fragrant white flowers (occasionally fading to creamy tones) in axillary clusters. Its pods are oblong and slightly inflated. Strictly endemic to the (primarily ), where it grows in subtropical laurel forests and rocky slopes, this species faces habitat threats from and development, considered threatened in regional assessments. Retama dasycarpa Coss., the least widespread, is an endemic Moroccan of undetermined typical height but similar to congeners (likely 2–4 m), with white or pale flowers and hairy pods (dasycarpa meaning "hairy fruit"). It exhibits habit with small leaves. Confined to Morocco's Mountains and semi-arid zones, it is adapted to rocky, soils, though specific data remain limited due to its rarity.

Infrageneric variation

Genetic diversity within the genus Retama varies among species, influenced by their geographic ranges and ecological niches. Retama raetam, with its extensive distribution across arid and semi-arid regions of the , , and the , displays relatively high ; a using random amplified polymorphic DNA (RAPD) markers on populations from southern found that 68% of the total variation occurred within populations, with significant differentiation among them (φPT = 0.316, p < 0.001), reflecting adaptation to diverse environmental conditions. In contrast, inter-simple sequence repeat (ISSR) analyses of Retama taxa in revealed moderate polymorphism, with 85 polymorphic bands out of 152 total, highlighting distinct genetic profiles between R. raetam and R. monosperma. Endemic species like Retama rhodorhizoides, confined to the , likely exhibit lower due to their restricted range, though dedicated studies are limited. Hybrids within Retama are rare, but potential has been noted in zones of , such as between R. monosperma and R. sphaerocarpa in southern and , where overlapping distributions may facilitate ; however, confirmed hybrid occurrences remain undocumented in molecular surveys. variation is prominent in R. raetam, which includes R. raetam subsp. raetam—a widespread Saharo-Arabian element distributed from to the —and subsp. gussonei, an endemic to southern and in , and subsp. bovei (endemic to ). These differ in geographic and subtle morphological traits, including pod dimensions and characteristics, with subsp. gussonei showing adaptations to Mediterranean coastal habitats. Molecular evidence from ISSR and RAPD markers supports clinal variation in Retama along arid environmental gradients, particularly in R. raetam, where genetic clusters correlate with habitat differences such as and levels in North African populations. Although nuclear ITS and plastid trnL-F regions have not been extensively applied to Retama, related studies suggest these markers could further resolve infrageneric relationships by detecting gradual shifts in drought-prone areas. Taxonomic implications from post-2020 emphasize the stability of core delimitations but highlight ongoing debates on infraspecific ranks; for instance, a 2023 revision in used molecular fingerprinting to affirm R. raetam and R. monosperma as distinct, while suggesting potential elevation of morphological forms to based on fruit and seed traits, potentially leading to splits in regional floras.

Human uses

Medicinal and pharmacological applications

Retama species have been employed in Mediterranean folk medicine for various therapeutic purposes, including as laxatives, diuretics, emetics, and vermifuges through decoctions prepared from leaves and branches. These preparations are also used topically to treat infections and orally to alleviate and joint pain. Historical dosages typically range from 5-10 g of dried material per day, often administered as infusions or decoctions, though exact quantities vary by region and preparation method. The phytochemical profile of Retama includes quinolizidine alkaloids such as , , and lupanine, primarily concentrated in roots and cladodes, alongside like and , and essential oils containing compounds such as and β-linalool in flowers. These bioactive constituents contribute to the plant's pharmacological potential, with alkaloids responsible for much of the therapeutic and toxic effects. Pharmacological studies demonstrate antioxidant activity in Retama extracts, with assay values ranging from 25 to over 1000 μg/mL, for example, 150 μg/mL for R. monosperma extracts, attributed to and content. Antimicrobial effects are evident against , with minimum inhibitory concentrations () of 0.256–1.25 mg/mL for various extracts. Anti-inflammatory properties involve inhibition of COX-2 and iNOS in models of acute , supporting traditional uses for . Modern research highlights antidiabetic potential through alpha-glucosidase inhibition by such as retamasin and erysubin from R. raetam, with values indicating competitive binding at the enzyme's . However, clinical trials remain limited, with most evidence derived from and animal studies. Toxicity concerns arise from , which can cause neurotoxic effects including CNS depression and , with oral LD50 values around 2000 mg/kg in rats for R. raetam extracts. Recent studies as of 2024 have further explored pharmacological activities. For instance, extracts of R. raetam leaves demonstrated properties and , promoting tissue regeneration and deposition in rat models. A 2023 review of R. monosperma confirmed its rich content of polyphenols, , and alkaloids, supporting , , and effects, with potential for treating skin diseases and . Additionally, the first and toxicological investigation of R. dasycarpa in 2024 revealed strong , , and antibacterial activities, highlighting its untapped potential as an endemic species.

Ornamental, ecological, and economic uses

Retama species, particularly Retama raetam and Retama sphaerocarpa, are valued in ornamental landscaping for their graceful, arching branches and profuse white or yellow blooms that appear in , adding aesthetic appeal to arid and Mediterranean gardens. These drought-tolerant shrubs are commonly planted in xeriscapes and as hedges, thriving in dry conditions with minimal irrigation once established, and are suitable for focal points or mixed borders in regions like and . In semi-arid landscapes, they provide year-round structure with their slender, green foliage and fragrant flowers, enhancing visual interest without requiring fertile soils. Ecologically, Retama plays a key role in stabilizing sandy dunes and slopes through its extensive root systems, which bind loose and prevent in arid environments. As nitrogen-fixing , species like R. raetam and R. sphaerocarpa form symbiotic relationships with bacteria, enriching nutrient-poor soils and supporting associated vegetation in systems. Additionally, Retama demonstrates potential for in heavy metal-contaminated sites, such as mine tailings rich in lead and , where it tolerates elevated metal levels and facilitates soil rehabilitation through nodulation in polluted conditions. Economically, branches of Retama serve as a reliable source of fuelwood, particularly for traditional ovens and heating in rural Mediterranean and North African communities, due to their dense wood and availability in scrublands. Fodder use is limited by the plant's toxicity—primarily from alkaloids in seeds and pods—but flowers and post-flowering foliage provide nutritious browse for resilient livestock like goats and dromedaries, supplementing diets in arid grazing areas. The nectar-rich flowers also support honey production, attracting bees and yielding monofloral honeys prized for their light color and floral notes in regions such as Andalusia and the Canary Islands. Cultivation of Retama for or ornamental purposes typically involves from , which require with hot water or nearly boiling water followed by soaking for 12-24 hours to achieve , or from semi-ripe cuttings in summer. Plants are spaced approximately 2-3 meters apart in projects to allow for root development and canopy spread, ensuring effective coverage while minimizing competition. In case studies from , Retama has been integral to efforts since the early 2000s, such as dune stabilization programs in Algeria's El-Mesrane region, where R. raetam plantings have enhanced , reduced sand mobility, and improved over decades compared to other shrubs like species. Similar initiatives in semi-arid utilize R. sphaerocarpa as a to reclaim degraded lands, promoting long-term recovery through enrichment and .

Cultural importance

Retama raetam, commonly known as the white or rotem, holds significant religious in traditions through its mention in the . In 1 Kings 19:4, the prophet travels into the wilderness and sits under a rotem in a state of deep isolation and despair, seeking shade from the harsh environment. This reference underscores the plant's role as a symbol of and , as an angel subsequently provides Elijah with sustenance while he rests beneath it. The rotem's ability to thrive in arid conditions further emphasizes themes of endurance and provision in biblical narratives. In North African and Middle Eastern , particularly among communities, R. raetam branches are used decoratively during cultural celebrations. Sprigs of the plant are fastened to the entrances of tents at weddings and other joyous occasions, serving as an expression of festivity and communal happiness. This practice highlights the plant's integration into social rituals, where its resilient growth in desert landscapes mirrors the enduring spirit of nomadic life. Retama species appear in and as emblems of the desert's harsh beauty and tenacity. Often portrayed as solitary figures enduring extreme , they evoke themes of steadfastness and in pre-Islamic and works, symbolizing the unyielding nature of the Arabian environment. In modern environmental writing, Retama's adaptations to are frequently cited to illustrate , reinforcing its metaphorical role in discussions of human perseverance amid adversity. In Mediterranean iconography, Retama is depicted as a hardy survivor, representing adaptation to challenging terrains in artistic representations of arid ecosystems. Its sparse, broom-like form appears in regional flora illustrations and cultural motifs, embodying endurance in landscapes from the to Iberia. In some arid regions, such as , it features prominently in biblical garden designs and national flora exhibits, symbolizing historical and environmental continuity. Contemporary cultural significance of Retama extends to eco-tourism and festivals in the , where species like Retama rhodorhizoides are celebrated as endemic symbols of the archipelago's unique . The plant's striking winter blossoms, known locally as "white magic," draw visitors to high-altitude trails in , integrating it into guided tours that highlight volcanic resilience and indigenous heritage. Annual events and botanical festivals often feature Retama, promoting awareness of its role in the islands' ecological and cultural identity.

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