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Phaseoleae

Phaseoleae is a diverse of flowering within the family , subfamily , encompassing approximately 90 genera and 1,570 species that are predominantly distributed in tropical and subtropical regions worldwide. These are typically herbs, vines, shrubs, or rarely trees, featuring pinnately trifoliolate leaves with pulvini, stipules, and stipels, as well as axillary racemose inflorescences bearing papilionaceous (butterfly-like) flowers with a 4- or 5-toothed and 2-valved, non-jointed containing strophiolate seeds. The is distinguished by its ability to form determinate "desmodioid" root nodules, which facilitate in with specific rhizobial . Phaseoleae holds significant economic and ecological importance, serving as a major source of food crops, , cover crops, green manures, ornamentals, medicines, and dyes. Notable genera include (soybean, G. max, a primary global oilseed and protein crop), Phaseolus (, P. vulgaris, and other New World beans), Vigna (cowpea, V. unguiculata, mung bean, V. radiata, and black gram, V. mungo, key pulses in ), and Cajanus (pigeon pea, C. cajan, a vital protein source in and ). These species contribute substantially to , , and sustainable farming practices due to their nitrogen-fixing capabilities, which enhance . Taxonomically, Phaseoleae is divided into seven subtribes: Cajaninae, Diocleinae, Erythrininae, Glycininae, Kennediinae, Ophrestiinae, and Phaseolinae, a classification that accounts for all recognized genera and reflects phylogenetic relationships based on morphological, chemical, and molecular data. The tribe's diversity is highest in the , with centers in the and , and ongoing research, including recent additions like the genus Weizhia (2023), addresses generic boundaries and evolutionary history through and nuclear phylogenies.

Description

General Morphology

Plants in the tribe Phaseoleae exhibit diverse growth habits, ranging from annual or perennial herbaceous vines and twining climbers to shrubs and, rarely, trees. Many species are scandent, utilizing tendrils derived from modified leaflets or stipules for support in climbing forms. Stems are typically herbaceous, varying from pubescent with hooked or appressed hairs to glabrous, and often exhibit longitudinal grooves or striations in twining species. Leaves are alternate and predominantly pinnately trifoliolate, with leaflets that are entire or occasionally lobed and sometimes gland-dotted; stipules and stipels are persistent or caducous, providing structural support at leaf bases. A defining vegetative feature of Phaseoleae is the presence of root nodules formed through with rhizobial bacteria, enabling . These nodules are of the determinate desmodioid type, characterized by spherical shape, prominent lenticels, and internal pink coloration due to , distinguishing them from indeterminate nodules in other legume tribes. This nodulation morphology is particularly adapted to the tribe's often tropical habitats and supports the nutritional demands of fast-growing climbers. Inflorescences in Phaseoleae are typically axillary and racemose or paniculate, bearing fascicles of flowers along the axis, though sometimes reduced to solitary blooms; bracts are small and early caducous, with nodes occasionally swollen. Flowers follow the papilionaceous (butterfly-like) structure typical of the subfamily , featuring a () petal, two petals, and a enclosing the stamens and , with diadelphous stamens (9+1) and a superior . Fruits are that are linear, falcate, oblong, moniliform, or irregularly inflated, usually dehiscent along both sutures with twisting valves, though some are indehiscent; they contain 2 to many seeds per pod, often partitioned between seeds. Seeds are typically reniform with a prominent hilum, strophiolate in some subtribes, and possess a hard testa that may be smooth, wrinkled, or reticulate.

Reproductive Features

The flowers of Phaseoleae exhibit the typical papilionoid structure characteristic of the subfamily , featuring zygomorphic symmetry with five sepals fused into a tubular , five distinct petals—including a broad standard (), two lateral wings, and a fused that encloses the stamens and —ten stamens arranged in a diadelphous (nine fused and one free), and a superior with a single carpel. These flowers often display nectar guides, such as prominent ridges on the petals that direct pollinators toward the nectar source, as observed in species like Vigna caracalla. Floral colors vary from white to shades of purple and vinaceous, enhancing visibility for pollinators. Pollination in Phaseoleae is primarily entomophilous, with bees serving as the main vectors through a "brush-type" mechanism where large bees, such as Bombus and Xylocopa species, trigger pollen release by manipulating the keel petals. Some species exhibit self-pollination, particularly in chasmogamous flowers that remain open but facilitate autogamy. Breeding systems within the tribe encompass a spectrum from predominantly outcrossing to selfing, with many taxa self-compatible yet reliant on pollinators for optimal gene flow; for instance, Phaseolus vulgaris is largely self-pollinating, while Phaseolus coccineus favors outcrossing. Cleistogamous flowers, which promote assured self-fertilization without pollinator access, occur in genera such as Vigna and Phaseolus acutifolius. Seed dispersal mechanisms in Phaseoleae predominantly involve ballistic dehiscence, where mature pods explosively split to propel seeds away from the parent plant, as seen in wild Phaseolus species where pod shattering facilitates escape from natural enemies. In other cases, dispersal is passive, relying on gravity or water currents to distribute seeds. Germination patterns vary, with both epigeal (cotyledons emerging above ground) and hypogeal (cotyledons remaining below ground) types represented; for example, Phaseolus vulgaris displays epigeal germination, whereas Phaseolus coccineus shows hypogeal germination. Tropical species often exhibit rapid early seedling growth, supported briefly by nitrogen-fixing root nodules that enhance nutrient acquisition post-germination.

Distribution and Ecology

Geographic Distribution

The tribe Phaseoleae displays a predominantly distribution, with native species spanning the , , , and , as well as extensions into subtropical and warm-temperate zones. This broad range reflects the tribe's adaptation to diverse tropical environments, encompassing approximately 84 genera and 1,500 species, many of which are concentrated in regions of high such as the seasonally dry tropics. Biogeographic patterns within the tribe show distinct centers of diversity, including high species richness in and the for New World lineages, and in eastern and for Old World groups. Key centers of origin align with major subtribes: the subtribe Phaseolinae, including economically important genera like Phaseolus, originated in the New World, with the highest diversity in Mesoamerica (particularly Mexico) and extending southward to northwestern Argentina and the northern Andes. In contrast, the subtribe Glycininae has Old World origins, centered in eastern Asia for annual species like Glycine soja and in Australia for numerous perennial taxa, contributing to elevated diversity in these regions. Other subtribes, such as those including Vigna and Dumasia, exhibit centers in tropical Africa and Asia, with Vigna showing additional diversification across Oceania and the Americas through historical dispersals. Human-mediated introductions via widespread cultivation have expanded Phaseoleae ranges beyond native areas, leading to naturalization in temperate , additional parts of , and even parts of outside core native zones. Patterns of endemism remain prominent, as evidenced by recent discoveries such as the monotypic genus Weizhia in eastern , highlighting ongoing revelations of localized diversity in subtropical .

Habitat and Adaptations

Species in the Phaseoleae predominantly occupy open woodlands, savannas, riverbanks, and disturbed areas, where they exhibit notable tolerance to seasonal . These habitats often feature variable moisture regimes, with many Phaseoleae thriving in seasonally dry tropical forests across , , and the . For instance, genera such as and Dolichos are commonly found in disturbed open savannas and wooded grasslands, benefiting from the periodic disturbance that maintains these ecosystems. Adaptations to in Phaseoleae include deep systems that access subsurface , succulent stems in certain genera like for storage, and efficient water-use strategies associated with . Herbaceous lineages within the , such as those in subtribe Phaseolinae, have diversified in response to Miocene , with shorter generation times facilitating rapid to dry conditions. Common bean () exemplifies this through its system, which penetrates deeply to mitigate stress, while ( unguiculata) employs similar architectures for resilience in semi-arid environments. Phaseoleae species prefer well-drained sandy or loamy soils, which support their growth in nutrient-variable environments, and their capacity for enables proliferation in poor soils. This symbiotic occurs via root nodules formed with specific rhizobial strains, such as and Ensifer for , and for , enhancing even in low-nitrogen substrates. Additionally, associations with arbuscular mycorrhizal fungi (AMF) improve uptake and , as seen in ( max), where AMF colonization boosts water absorption and overall resilience. Ecological interactions in Phaseoleae include vulnerability to pests such as beetles, including the Mexican bean beetle (Epilachna varivestis) on Phaseolus crops and seed beetles like on stored , which can significantly impact wild populations in disturbed habitats. Regarding , many Phaseoleae are thermophilic, adapted to warmer tropical and subtropical conditions, though some remain frost-sensitive, limiting their range in temperate zones. In habitats, species exhibit fire responses such as seed scarification for and resprouting from underground organs, promoting post-fire recovery in fire-prone ecosystems.

Taxonomy and Classification

Historical Development

The tribe Phaseoleae was first recognized as a distinct group within the subfamily Papilionoideae by in , based on morphological characteristics such as the structure of the flowers and fruits, particularly the dehiscent pods that often twist elastically upon opening. Bentham's initial delineation emphasized the papilionaceous and the linear, multi-seeded , distinguishing Phaseoleae from other tribes like Dalbergieae and . This classification laid the foundation for subsequent taxonomic work, highlighting the tribe's tropical affinity and economic potential through genera like . In Bentham's comprehensive treatment in Genera Plantarum (1865), Phaseoleae was formalized as one of 11 tribes in Papilionoideae, with an initial subtribal framework that included groups like Phaseolinae and Erythrininae, delimited primarily by pod dehiscence patterns, flower symmetry, and morphology. This scheme recognized around 50-60 genera, focusing on vegetative and reproductive traits such as the twisting of pod valves and the arrangement of floral bracts, which were seen as synapomorphies for the tribe. Over the late 19th and early 20th centuries, refinements incorporated new species descriptions, but the core morphology-based subtribes persisted, with additions like Diocleinae based on keel petal fusion and seed development. Twentieth-century classifications introduced biochemical markers alongside morphology, notably through J.A. Lackey's work in the 1970s, which emphasized the distribution of the non-protein amino acid as a taxonomic indicator, particularly its prevalence in subtribes like Diocleinae and Kennediinae. Lackey's 1977 revision proposed seven subtribes, integrating presence with traits like attachment and structure to resolve ambiguities in Bentham's system, while accommodating newly described genera such as Neonotonia. By Lackey's 1981 account in Advances in Legume , the tribe encompassed approximately 84 genera and 1,480 , reflecting ongoing shifts from morphological re-evaluations and the exclusion of atypical elements like certain species. Pre-molecular taxonomy in the 1980s and 1990s relied heavily on fruit and floral , such as pod twisting and lip development, leading to the recognition of about 80 genera by the early 1990s through works like those of Polhill (1994), which maintained Lackey's subtribal structure while noting inconsistencies in genera like Psophocarpus. The transition to molecular approaches began in the early 2000s, with studies using rbcL sequences revealing in Phaseoleae, as subtribes like Ophrestiinae and Diocleinae nested within Millettieae, prompting mergers of adjacent tribes and a reevaluation of boundaries. These findings, exemplified by Kajita et al. (2001), underscored the limitations of morphology alone and set the stage for broader phylogenetic revisions.

Current Subtribes

The modern subtribal classification of the tribe Phaseoleae follows the framework established by Lackey in 1977, which recognizes seven principal subtribes along with a few genera of uncertain placement (). This system integrates morphological traits such as structure, floral features, and characteristics to delineate groups, and it has been largely upheld in subsequent taxonomic revisions, though molecular have prompted minor adjustments. The tribe encompasses approximately 84 genera and 1,500 as of 2024, reflecting updates that incorporate newly described taxa like Weizhia (placed in Glycininae) based on combined morphological and phylogenetic evidence. Cajaninae includes about 5 genera, such as (the pigeonpea group), and is distinguished by the presence of vesicular glands on leaves and stems, bulbous-based hairs, and often dehiscent pods with canavanine distribution patterns. Diocleinae contains approximately 15 genera, including Dioclea and Canavalia (jack beans), characterized by arillate seeds, often with a prominent strophiole, and vines or lianas with pseudoracemose inflorescences; canavanine is commonly present. Erythrininae features genera like (coral trees), with wing petals longer than the keel and often scarlet flowers adapted for bird pollination, though its boundaries have been refined post-2013 phylogenetic updates. Glycininae encompasses around 10 genera, such as (soybeans) and the recently added Weizhia, notable for pseudoracemose inflorescences where flowers are borne on leafy shoots, lomentaceous or indehiscent pods, and frequent absence of . Kennediinae consists of about 8 genera, like Kennedia, with prominent arils, absence of bracteoles, and climbing habits suited to temperate to subtropical environments. Ophrestiinae is a small subtribe with 1–2 genera, including Ophrestia, defined by unique pod dehiscence mechanisms and limited distribution, often in tropical regions. Phaseolinae, the largest subtribe with approximately 20 genera such as Phaseolus (common beans) and Vigna (cowpeas), is identified by valvate sepals, explosive dehiscing pods that aid seed dispersal, and bearded styles; some genera like Rhynchosia have seen synonymy reductions through taxonomic revisions. A handful of genera remain incertae sedis pending further molecular resolution.

Phylogeny and Evolution

Molecular Phylogenetics

Molecular phylogenetic studies of the Phaseoleae tribe have primarily relied on DNA sequence data from and nuclear ribosomal s to resolve relationships within the broader Millettioid/Phaseoloid (MP) clade of Papilionoideae. Early analyses, such as Wojciechowski et al. (2004), utilized the plastid matK across 330 sequences from 235 genera, revealing the MP clade as a well-supported monophyletic group comprising approximately 300 genera and over 6000 species, with Phaseoleae forming one of two major subclades alongside Millettieae. This study highlighted the of traditional Phaseoleae, as several genera traditionally placed therein nested outside the core group. Subsequent multi-gene approaches expanded on these findings, incorporating both and markers for finer resolution. Stefanović et al. (2009) analyzed sequences from eight regions (rbcL, atpB, trnK/matK, trnL-trnF, rpl32-trnL, ycf3-trnS, psbA-trnH, and ndhF) across 115 phaseoloid taxa, confirming the position of Phaseoleae within the NPAAA clade (a basal papilionoid assemblage characterized by the absence of the typical 50 kb inversion in the large single-copy region of the genome). Their phylogeny demonstrated that Phaseoleae is paraphyletic with respect to Desmodieae and Psoraleeae, which are nested within it, while the core Phaseoleae—encompassing subtribes like Phaseolinae and Glycininae—emerged as monophyletic with strong bootstrap support (>90%) but exhibited basal divergences among early-branching lineages. markers, including the (ITS) and external transcribed spacer (ETS) of , have been integrated in complementary studies to address cytonuclear discordance and provide additional evidence for these relationships. Recent advances in high-throughput sequencing have shifted toward whole plastome analyses, offering greater phylogenetic signal due to increased locus coverage. For instance, Wang et al. (2024) sequenced and compared the complete plastome of Amphicarpaea edgeworthii (hog-peanut, Glycininae) with 34 other Phaseoleae taxa, reconstructing a phylogeny that reinforces the of core Phaseoleae, with posterior probabilities exceeding 0.95 at key nodes. These findings support proposed taxonomic revisions, such as the merger of certain Millettieae genera (e.g., those in the Ophrestiinae subtribe) into an expanded Phaseoleae, based on shared synapomorphies and phylogenetic nesting. Recent phylogenomic studies, including those on Asian s.l., have further refined generic boundaries within Phaseoleae, confirming robust support for major subtribes. Within core Phaseoleae, plastome data affirm robust boundaries for major subtribes: Phaseolinae (including economically vital genera like and ) forms a well-supported sister to Glycininae (e.g., and Amphicarpaea), with divergence estimates aligning with Oligocene-Miocene radiations.

Diversification Patterns

The tribe Phaseoleae, part of the subfamily , originated through a divergence event in the late Eocene to early , with molecular estimates placing the stem age at approximately 39.5 million years ago (Ma) and the crown age at around 28.6 Ma. Early lineages likely emerged in an African-Asian cradle, centered in , where initial diversification occurred amid climatic transitions. Following the breakup of in the , subsequent radiations into the took place during the , facilitated by intercontinental dispersals and the opening of new ecological niches in the Neotropics. Major diversification events in Phaseoleae align with environmental changes, including expansions linked to the global evolution of grasslands around 18–8 Ma. This period saw rapid , particularly in the subtribe Phaseolinae, estimated at 10–15 Ma, driven by the proliferation of open habitats and herbaceous growth forms that allowed exploitation of disturbed environments. These events involved shifts in net diversification rates, with elevated speciation following the Mid-Miocene Climatic Optimum, as ancestral Asian-African lineages dispersed widely. Key drivers of speciation included climate shifts toward during the Oligocene-Miocene transition, which promoted adaptations to seasonally dry forests and grasslands, and biogeographic barriers such as the uplift of the starting in the mid-Miocene, which isolated populations and spurred adaptive radiations in . also played a significant role in certain genera, notably , where reticulate evolution through allopolyploid events enhanced genetic diversity and facilitated niche expansion in and beyond. Some incertae sedis taxa within Phaseoleae represent evolutionary relics, with observed in clades attributable to incomplete lineage sorting from ancient polymorphisms. The fossil record of Phaseoleae remains limited, with few unambiguous legume fossils directly attributable to the tribe, though Miocene pod impressions from , such as those of the extinct genus Podocarpium, provide evidence of early presence in and support timelines of Asian diversification. These fossils, dating to around 13–15 Ma, highlight the tribe's adaptation to emerging Miocene biomes but underscore gaps in understanding extinction dynamics, as many relictual forms may have succumbed to climatic volatility.

Diversity and Genera

Major Genera

The genus Phaseolus includes approximately 87 , all native to the from the to northern . These species are predominantly herbaceous to woody annual or perennial vines that twine, featuring characteristic phaseoloid pods that are linear and often dehiscent. The genus encompasses economically vital members such as the common bean (P. vulgaris), a key domesticated . Vigna is one of the most species-rich genera in Phaseoleae, with 107 accepted species distributed pantropically across , , , and the . Plants in this genus are typically prostrate, trailing, or climbing herbs and subshrubs, often producing small, rounded seeds adapted for wide dispersal. Notable examples include the (V. radiata), an important Asian crop. The genus Glycine contains 28 accepted species, primarily endemic to Australasia including Australia, New Guinea, and parts of Southeast Asia. These are mostly scrambling or twining perennials with pseudoracemose inflorescences, and polyploidy is prevalent, contributing to genomic complexity in the group. The soybean (G. max) represents a cornerstone domesticated species within the genus. Other prominent genera include , which comprises 34 species mainly in the Old World tropics from to and , often as erect shrubs or climbers with indehiscent pods; it features the (C. cajan) as its flagship species. Macroptilium has 22 species native to tropical and subtropical , characterized by trailing or climbing habits and used in forage; siratro (M. atropurpureum) is a key representative. Lablab is monotypic, consisting solely of (hyacinth bean), a robust climbing perennial native to tropical and southern with purple flowers and edible pods. A recently described genus is Weizhia, established in 2023 as a monotypic endemic to eastern China in Zhejiang Province, with the single species W. pentaphylla distinguished by its pinnately 5-foliolate leaves, spurred wing petals, and placement in subtribe Glycininae based on molecular evidence.

Species Diversity

The tribe Phaseoleae encompasses approximately 84 genera and 1,500 species, making it one of the largest tribes within the Fabaceae family. Estimates suggest a range of 1,500–1,600 species across 84–90 genera, reflecting ongoing taxonomic revisions and discoveries. Diversity is concentrated in certain subtribes. Diversity hotspots for Phaseoleae are prominent in tropical and subtropical regions, particularly where Phaseolus species dominate, with high Vigna representation, and hosting diverse genera across varied habitats. These areas exhibit elevated due to favorable climatic conditions and historical biogeographic factors. Patterns of endemism in Phaseoleae are notable in insular and montane environments, with high levels observed in islands such as , particularly within Clitoriinae, and in isolated mountain ranges where narrow-range species predominate. Introduced species also pose risks as invasives in non-native regions, complicating management efforts. Since 2020, taxonomic research has described 5–10 new species within Phaseoleae, including Apios chindiana from in 2025, often from understudied montane and habitats. A 2019 revision of Sphenostylis in also highlighted potential for further additions through refined .

Economic and Cultural Importance

Agricultural Crops

The Phaseoleae tribe encompasses several major agricultural crops that play pivotal roles in global , providing essential proteins, oils, and . Soybean (Glycine max) stands as the most prominent, with global production reaching approximately 424 million metric tons as of 2024/25, primarily serving as a source of high-quality protein for and vegetable oil for human consumption and industrial uses. Common (Phaseolus vulgaris) follows as a vital staple, yielding around 28.9 million metric tons as of 2024 and forming a dietary cornerstone in and , where it contributes significantly to protein intake in resource-limited diets. Other key Phaseoleae crops include (Vigna radiata), a central to Asian with global output of about 10.6 million tons from 8 million hectares as of 2022, valued for its quick maturation and nutritional profile in soups, sprouts, and snacks. (Cajanus cajan) offers drought tolerance, producing roughly 4.8 million tons worldwide as of 2022, particularly in semi-arid regions of and , where it supports systems for and . (Vigna unguiculata) yields approximately 9 million tons as of 2021, with over 95% from , serving dual purposes as a for human consumption and for livestock in . Domestication of these crops occurred independently around 8,000–10,000 years ago, with common bean originating in , soybean in , and Vigna species like and in and , marking early agricultural innovations that enhanced and through . Phaseoleae crops are major contributors to global production, underscoring their economic scale despite challenges such as yield losses from diseases, including (Uromyces appendiculatus) in common bean, which can reduce harvests by up to 85% in susceptible varieties. Recent breeding advances, including varieties developed via , have improved pest resistance and climate adaptation, enabling higher yields under and stress while preserving from wild ancestors.

Other Uses and Conservation

Species in the Phaseoleae tribe have applications beyond major agricultural crops, including medicinal, ornamental, and ecological roles. Several genera exhibit ethnomedicinal properties, with extracts used traditionally for treating various ailments. For instance, species of Rhynchosia (Cajaninae subtribe) are employed in and Asian as pain relievers, aphrodisiacs, and remedies for conditions like and inflammation, supported by phytochemical analyses revealing with , , and anticancer activities. In Argentine-Chilean , Phaseolus vulgaris, Phaseolus coccineus, and Phaseolus lunatus are gathered from wild habitats for medicinal purposes, often in infusions to address digestive and respiratory issues, based on ethnobotanical surveys of 63 species. Additionally, Clitoria ternatea (Clitoriinae subtribe) is valued in Ayurvedic medicine for its and memory-enhancing effects, attributed to bioactive compounds like ternatins. Ornamental uses are prominent in certain genera due to their attractive flowers and growth habits. species, known as coral trees, are widely cultivated for their vivid red blooms and are used in landscaping across tropical regions, with noted for its hardiness and showy inflorescences in subtropical gardens. Similarly, , or butterfly pea, is grown as an ornamental climber for its striking blue flowers, often in revegetation projects and home gardens, requiring minimal maintenance once established. Ecologically, Phaseoleae species contribute to through symbiotic with , forming determinate nodules that enhance nutrient availability in agroecosystems. This trait is well-documented in (common bean), where rates vary with environmental factors but support sustainable cropping by reducing fertilizer needs. Conservation efforts for Phaseoleae are critical due to habitat loss and narrow distributions affecting several endemics. Three Costa Rican Phaseolus species—P. albicarminus, P. angucianae, and P. hygrophilus—are assessed as Endangered (EN) under IUCN criteria, with extents of occurrence below 150 km² and threats from deforestation for agriculture and grazing, prompting calls for protected area inclusion. In South Africa, Rhynchosia ngwenyii is Endangered owing to its restricted range in KwaZulu-Natal and Mpumalanga, vulnerable to habitat fragmentation. The newly described genus Weizhia from East China, comprising a single species, is Critically Endangered (CR) due to urbanization pressures on its coastal habitat, highlighting the need for immediate ex situ conservation. Overall, while the tribe shows lower endemism in regions like southern Africa (66 species with residual endemism of -77), targeted protections are essential to preserve biodiversity amid global threats.