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Cajanus

Cajanus is a of flowering plants in the legume family , subfamily Papilionoideae, comprising 34 primarily distributed across , , and . The genus is characterized by its woody shrubs or small trees, often with trifoliate leaves and yellow flowers, and plays a significant role in due to its nitrogen-fixing capabilities. The only cultivated , Cajanus cajan (), is a short-lived shrub growing up to 4 meters tall, native to the but now widely grown in semi-arid worldwide. Pigeon pea serves as a major protein source for over 1 billion people in developing regions, with its seeds containing 20–22% protein and used in various culinary forms such as or . Beyond , the plant provides for livestock, for soil fertility, and wood for fuel, while its deep system aids in and in rain-fed cropping systems. Domesticated around 3,000–3,500 years ago in , C. cajan is intercropped with cereals and other crops, contributing to sustainable farming in over 80 countries, particularly in (where accounts for approximately 75% of global production as of 2023) and sub-Saharan Africa. The wild relatives of Cajanus form three gene pools: the primary pool including C. cajanifolius, from which the crop likely originated; a secondary pool of 10 cross-compatible ; and a tertiary pool of the remaining non-crossable , which offer potential for breeding traits like disease resistance. Morphologically, species in the genus feature pinnately trifoliate leaves, zygomorphic flowers in racemes, and pods with 2–9 seeds, adapting well to diverse habitats from to 3,000 meters altitude.

Taxonomy

Etymology

The genus name Cajanus derives from the word "kacang" (formerly spelled "katjang"), meaning "" or "bean," in reference to the plant's pods. This reflects its significance as a pod-bearing legume in tropical regions. formally proposed the genus as Cajan in his 1763 work Familles des Plantes, separating it from the broader genus based on observations of cultivated forms from tropical regions, including . However, Adanson's name was later considered illegitimate due to nomenclatural issues and lack of a designated . In the early 19th century, established the modern Cajanus in 1813, formally validating it with the description of two , C. flavus and C. bicolor, both attributed to origins and encompassing cultivated variants. De Candolle further refined the in his 1825 Prodromus Systematis Naturalis Regni Vegetabilis, incorporating Cytisus cajan L. (from Carl Linnaeus's 1753 ) as a and emphasizing morphological distinctions such as structure and habit. This revision solidified Cajanus as a distinct entity within the Leguminosae (now ), addressing earlier confusions with related genera. Historical synonyms persisted, including cajan L. and Cajanus indicus Spreng., highlighting the plant's prior misclassifications under broom-like taxa. The nomenclature was stabilized in the through international congresses; at the 1910 Brussels Botanical Congress, Cajanus . was conserved over Adanson's earlier Cajan, ensuring its priority and widespread acceptance. This timeline—from Linnaean placement in 1753, Adanson's proposal in 1763, de Candolle's foundational work in 1813–1825, to modern conservation—marks the evolution of the genus name amid expanding knowledge of its Asian origins and global distribution. Today, Cajanus is firmly recognized in subtribe Cajaninae of tribe in subfamily , family , encompassing 34 to 37 species as of 2025, with the C. cajan.

Classification

Cajanus belongs to the kingdom Plantae, phylum Tracheophyta, class Magnoliopsida, order , family , subfamily , tribe , and subtribe Cajaninae. This placement reflects its position among the , characterized by typical fabaceous features such as compound leaves and nitrogen-fixing root nodules. The underwent significant taxonomic revision in 1985 by L.J.G. van der Maesen, who recognized in Cajanus, incorporating several taxa previously assigned to the closely related genus Atylosia and describing three new species from . Subsequent assessments have estimated the total number of species between and 37, primarily distributed in tropical regions, though exact counts vary due to ongoing taxonomic refinements; as of , authoritative sources report 34 accepted species. Phylogenetic analyses based on molecular data, including nuclear ribosomal (ITS) sequences, have demonstrated that Cajanus forms a monophyletic group and exhibits close evolutionary relationships with genera such as Rhynchosia and Atylosia within subtribe Cajaninae. These studies, conducted in the and early , utilized markers like restriction fragment length polymorphisms (RFLPs) and sequence data to resolve intergeneric affinities, supporting the consolidation of related taxa under a unified phylogenetic framework.

Description

Morphology

Plants in the genus Cajanus exhibit diverse growth habits, ranging from erect shrubs and subshrubs to or trailing vines, typically reaching heights or lengths of up to 4 meters. The stems are often pubescent and can be spreading to erect in shrubby species or slender and twining in climbing forms, arising from a woody or deep system. Leaves are alternate and odd-pinnately compound, usually trifoliolate with three leaflets; the leaflets are elliptic to lanceolate, measuring 2-10 cm in length and 1.5-4 cm in width, often with pubescence and glandular dots on the surfaces. Stipules and stipels are present, and the petiole ranges from 1-5.5 cm long. Inflorescences are terminal or axillary racemes (sometimes appearing as pseudo-racemes in panicle form), bearing papilionaceous flowers that are typically yellow to orange, with a campanulate calyx, broadly ovate standard (1.2-1.7 cm diameter), oblong wings, and a spirally incurved keel; the flowers measure up to 3 cm and feature diadelphous stamens with uniform anthers and a glabrous, upcurved style ending in a capitate stigma. The fruits are linear, straight or slightly falcate pods, 2-10 cm long and 5-14 mm wide, dehiscent, and pubescent or glandular, containing 2-9 reniform with a strophiole; the is sessile with few to many ovules. Morphological variations are notable across species, such as the erect, woody of C. cajan contrasting with the twining, stems of C. scarabaeoides, which forms mats up to 2 m long and has smaller, pubescent trifoliolate leaves. Similarly, C. cajanifolius, a close relative of the cultivated , shares the erect form but displays subtle differences in leaflet size and pubescence compared to C. cajan.

Reproduction

Cajanus species primarily reproduce sexually through hermaphroditic flowers that are zygomorphic and typically yellow, with promoted via mechanisms such as weak , protogyny, and occasional male sterility as observed in the cultivated species C. cajan. In the cultivated species C. cajan, natural rates vary widely from 5% to 70%, influenced by environmental factors, insect pollinators, and , with an average of around 20–30% under typical conditions. occurs mainly through cross-pollination, though is possible; anthers dehisce a day before flower opening, with peak between 09:00 and 10:00 hours, leading to fertilization on the same day. Seed development in C. cajan results in pods containing 2–9 seeds, which reach physiological maturity about 30 days after and are harvestable at 40 days. These exhibit physical due to a hard impermeable seed coat, which imposes a dormancy period that can last several months post-harvest; rates remain low (around 10–11% in untreated wild relatives) without treatments like or hot water soaking, but can exceed 94–100% after such interventions in accessions. Optimal occurs at temperatures of 29–36°C, with the testa splitting on day 2, emergence on day 3, and development shortly thereafter; seeds are and can be stored long-term under frozen conditions. Asexual reproduction is rare across the but has been observed through vegetative propagation in cultivated C. cajan, particularly via stem cuttings, though success rates are low and not commonly practiced. Species in the Cajanus display life cycles ranging from annuals to or short-lived , with C. cajan typically functioning as a short-lived (up to 5 years) but often managed as an . Flowering is triggered by short day lengths in tropical environments, as C. cajan is a quantitative short-day sensitive to photoperiods below approximately 12–12.5 hours, with short-season varieties initiating blooms in 60–80 days and longer-duration types requiring 180–250 days. Seedlings emerge 2–3 weeks after , with initial growth slow until the establishes.

Distribution and habitat

Native range

The genus Cajanus is primarily native to tropical regions of , , and , encompassing approximately 32 wild species adapted to diverse environments in these areas. In tropical , the native range includes —such as , , , , , and —along with and nearby islands like and the , where species like C. kerstingii occur endemically in . Across Asia, the distribution spans the , Indochina (including , , and ), southern (notably ), and (encompassing the , , the , , and extending to the of ), with the serving as a major center of diversity hosting 17 species, many of which are endemic, particularly in regions like the , , and Nilgiri Hills. In Australasia, native populations are concentrated in (across , the , and ) and , where 13 species are endemic to and one extends to . Wild Cajanus species inhabit seasonally dry tropical forests, open woodlands, grasslands, savannas, scrub vegetation, and rocky outcrops such as hills, typically in semi-arid to semi-humid tropical climates. Their altitudinal distribution ranges from to 2,700 meters.

Introduced ranges

Cajanus species, particularly C. cajan, were introduced to regions beyond their native ranges in , , and through ancient and colonial routes, with significant expansions occurring since the . The plant reached tropical around 2000 BCE via early migrations and , establishing secondary centers of diversity there. By the 17th century, European colonialism and the transatlantic slave facilitated its spread from African ports in and to the tropical , including , the (such as and the ), and , where it was transported by enslaved people as a resilient source. As of 2023, C. cajan is cultivated in over 80 countries across tropical and subtropical zones, with major production in (approximately 85% of global output), (approximately 12%), and the (approximately 3%). Beyond formal agriculture, naturalized populations of C. cajan have established in disturbed habitats such as roadsides, fallow lands, and forest edges in the Neotropics and Pacific islands like , where escapes from cultivation have persisted since introductions in the 19th century. The successful establishment of Cajanus in introduced ranges is aided by its symbiotic nitrogen-fixing capabilities with bacteria, which enhance in nutrient-poor, acidic, or degraded environments, allowing persistence in marginal lands without heavy inputs. This trait, combined with and adaptability to a wide pH range (4.5–8.0), has enabled self-sustaining populations and reduced reliance on intensive management in new habitats.

Ecology

Pollination and dispersal

In natural populations of Cajanus, is predominantly facilitated by , with of the genus Xylocopa serving as primary pollinators due to their ability to trip the flowers and effect transfer from the anthers to the . Other bees, including honey bees (Apis mellifera) and solitary bees such as Megachile species, also contribute significantly to transfer, visiting flowers for and . Moths occasionally participate in cross- by accessing the floral rewards. The structure, featuring a that encloses the reproductive organs, promotes this insect-mediated by requiring physical agitation to release . In cultivated C. cajan, self-pollination occurs through cleistogamous flowers, but natural outcrossing rates range from 20% to 70%, influenced by insect activity and environmental factors. Wild Cajanus species also rely on insect-aided cross-pollination. Pollination efficiency varies by vector; for instance, Xylocopa olivacea foraging on C. cajan flowers at rates of approximately 12 flowers per minute enhances fruit set by up to 22% and seed production per pod by 35%, demonstrating the critical role of these insects in reproductive success. Seed dispersal in natural Cajanus populations occurs mainly through autochory, where dry pods dehisce, releasing seeds that fall nearby via gravity to establish seedlings in close proximity to the parent plant. Long-distance dispersal is primarily mediated by human activities, such as through trade and agriculture, which have facilitated the genus's introduction beyond its native range. These mechanisms ensure seed persistence in diverse habitats, with hard seed coats in wild species enhancing survival against predation and environmental stresses during dispersal.

Interactions with animals

Cajanus species, particularly C. cajan, serve as hosts for various insect herbivores, including larvae of Lepidoptera moths. For instance, the ghost moth Endoclita malabaricus feeds on C. cajan foliage during its larval stage, contributing to defoliation in native habitats. Additionally, pod borers such as Helicoverpa armigera and Maruca vitrata target developing pods, causing significant damage in cultivated fields by boring into seeds and reducing yield. Goats occasionally browse on C. cajan leaves and stems, especially in mixed agroforestry systems where the plant is used as fodder, though this interaction is typically limited to avoid overgrazing. Wild Cajanus species exhibit defensive traits that deter herbivores, such as elevated levels of and isoflavonoids, which impart bitterness and reduce . In C. scarabaeoides, a close relative of the cultivated , these secondary metabolites correlate with resistance to pod borers, slowing larval development and inhibiting feeding compared to domesticated varieties that have lost some of these compounds during breeding. This biochemical defense highlights the genus's adaptation to biotic pressures in natural ecosystems. In contrast to these antagonistic animal interactions, Cajanus engages in mutualistic with rhizobial like species, which fix atmospheric nitrogen in root nodules to enhance plant growth and in natural habitats, without involving . Within agroecosystems, C. cajan crops indirectly support populations by providing resources that sustain communities, though these benefits are secondary to herbivory pressures.

Cultivation and uses

Domestication and agriculture

The domestication of Cajanus cajan, commonly known as pigeon pea, occurred in India approximately 3,500 years ago, originating from its wild progenitor Cajanus cajanifolius. Archaeological evidence from sites in the Indus Valley and Orissa supports early cultivation dating to the mid-second millennium BC, marking it as one of the oldest domesticated pulses in South Asia. From its center of origin in central and eastern India, the crop spread through ancient trade routes to Southeast Asia, East Africa, and eventually the Americas during colonial periods, adapting to diverse tropical environments. In modern agriculture, pigeon pea is widely cultivated as a rainfed in tropical and subtropical regions, often intercropped with cereals like or to improve , reduce , and stabilize yields in low-input systems. It performs best in well-drained loamy with a of 5.0 to 7.0, tolerating through its deep system but requiring moderate rainfall of 600–1,000 mm annually for optimal growth. Global average yields for dry seeds range from 700 to 1,000 kg/ha (as of 2021), though intercropping systems often achieve lower outputs of 400–500 kg/ha due to competition, while improved practices can exceed 2,000 kg/ha. Breeding efforts for intensified in the late , led by institutions such as the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), focusing on enhancing , resistance to like pod borers, and overall yield stability through hybridization with wild relatives and genomic selection. These programs have developed short-duration varieties that mature in 120–180 days, enabling integration into intensive cropping rotations and addressing climate variability in marginal lands. As of 2025, ICRISAT has developed the world's first extreme heat-tolerant variety, ICPV 25444, using speed-breeding protocols, maturing in 125 days under 45°C conditions, along with new genes for disease resistance and high-protein cultivars.

Food and medicinal applications

Pigeon pea (Cajanus cajan) seeds are a in many tropical regions, particularly in and , where they are consumed for their high nutritional value. The mature seeds provide approximately 18-25% protein on a dry weight basis, making them a valuable plant-based protein source comparable to other . This protein is rich in essential such as (70 mg/g protein) and (71 mg/g protein), though it is relatively deficient in sulfur-containing amino acids like . In culinary applications, the dehulled and split seeds, known as dhal or , are widely used in n cuisines for preparing dals, curries, and rice dishes, often complementing cereal-based diets to enhance protein intake. In African diets, whole mature seeds are boiled or germinated for porridges and stews, while green seeds and pods serve as in salads and soups. In , an estimated 65% of produced pigeon peas are consumed by farmers themselves. The seeds are also ground into flour for flatbreads or blended into fermented products to improve digestibility and nutritional bioavailability. Beyond nutrition, C. cajan has significant medicinal applications rooted in traditional systems across and . In folk medicine, the leaves and roots are used to treat ailments such as —often chewed or applied as a paste for and sore gums—and , with decoctions believed to regulate blood sugar levels, as practiced by communities in , , and . Experimental studies support these uses; for instance, methanolic extracts of the leaves demonstrated antidiabetic effects in alloxan-induced diabetic rats by significantly reducing fasting blood glucose levels in a dose-dependent manner (200-600 mg/kg), with peak effects observed 4-6 hours post-administration. These effects are attributed to bioactive , including cajanin (an found in seeds and stems), , and orientin, which inhibit α-glucosidase enzymes and enhance insulin sensitivity. Clinical and research from the 2010s further highlights the properties of C. cajan extracts, which scavenge free radicals and boost endogenous enzymes like . Ethanol extracts of seeds exhibited concentration-dependent activity (25-450 µg/mL) in FRAP assays, correlating with high and contents that mitigate linked to and . Root extracts similarly showed strong and ABTS radical scavenging, supporting their traditional use in managing chronic conditions. Among other Cajanus species, C. scarabaeoides has minor applications, primarily as for in tropical regions, where its stems and crushed seeds provide nutritive feed that also helps reduce in .

Diversity

Number and distribution of species

The genus Cajanus comprises 32–37 accepted , reflecting ongoing taxonomic refinements. Approximately 17 species are endemic to the , 13 to , and the remainder occur in and other parts of , with one species (C. kerstingii) restricted to western . These species are predominantly distributed in tropical regions of the , with major centers of diversity in seasonally dry habitats such as savannas and deciduous forests; species outside the are exceedingly rare and typically represent introductions. Taxonomic challenges in the genus include historical mergers of related genera like Atylosia into Cajanus based on morphological and cytological evidence, with recent molecular phylogenetic studies in the 2020s contributing to synonymizations and clearer delineation of species boundaries.

Notable species

The most prominent species in the genus Cajanus is C. cajan (L.) Millsp., known as pigeon pea, the sole cultivated member of the genus and a staple food legume in tropical and subtropical agriculture. This short-lived perennial shrub is valued for its protein-rich seeds, which supply essential amino acids like methionine and lysine, supporting nutrition in regions with limited animal protein access. Beyond food, C. cajan is utilized as a green manure to enhance soil nitrogen, a forage crop for livestock, and a source of fuelwood, with global production exceeding 5 million metric tons annually as of 2022, primarily in India. Several wild relatives stand out for their contributions to pigeon pea breeding and genetic conservation, given the genus comprises about 34 , mostly uncultivated. C. scarabaeoides (L.) Thouars, native to , , and , is the nearest wild progenitor and has been hybridized with C. cajan to transfer traits such as for hybrid seed production and resistance to pests like pod borers. C. cajanifolius (Haines) Maesen, endemic to , serves as a key source for genes, while C. platycarpus (Benth.) Maesen, found in , provides alleles for early maturity, aiding to shorter growing seasons. These , prioritized for due to habitat threats, underscore the genus's role in sustainable crop improvement.

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