Diversity of thrips (order: Thysanoptera) in leguminous crops: A comprehensive review

Introduction

Leguminous crops (Family: Fabaceae) are important food sources with high protein content and are adaptable to diverse agroclimatic conditions. It provides essential dietary fibres and protein for human and livestock consumption (Singh et al., 2003) [80] , while contributing to atmospheric nitrogen fixation through root nodules, making them an integral part of sustainable agricultural production systems (Kolawole et al., 2000; Giller, 2001; Zhu et al., 2005) [21,39,98] . It contributes to the supply of lysine-rich proteins, which are deficient in cereal-based diets (Ufaz and Galili, 2008) [90] . It helps alleviate poverty and micronutrient deficiencies among millions of people around the world. Major legume crops such as groundnut (Arachis hypogaea), soybean (Glycine max), pigeon pea (Cajanus cajan), common bean (Phaseolus vulgaris), cowpea (Vigna unguiculata), chickpea (Cicer arietinum) and lentil (Lens culinaris) are grown in tropical, subtropical and temperate regions. Pulses occupy 90-96 million hectares, producing 87.7 million tonnes globally (FAOSTAT, 2023) [18] . However, insect pests are among the major biotic stresses, particularly thrips, which are affecting legumes and hindering the attainment of maximum yield potential. These destructive pests belong to the order Thysanoptera, and the major agriculturally important species belong to the family Thripidae, suborder Terebrantia. Major morphological characters include asymmetrical mouthparts, with the right mandible absent, and fringed wings, with complex life cycles that include two larval instars and two pupal stages (prepupa and pupa) (Kumar and Omkar, 2021) [42,43] . The shorter lifecycle leads to rapid reproduction rates, producing several generations that contribute to their explosive population dynamics and invasiveness (Lewis, 1973) [44,45] Thrips cause direct feeding damage, leading to leaf distortion, flower abortion, and bronzing of pods, resulting in yield losses of 20-80% (Singh and Allen, 1980) [81] . Flower bud abortion is a major constraint affecting pod yield. Their role as vectors of tospoviruses such as Tomato Spotted Wilt Virus (TSWV) is equally significant, causing severe yield losses in crops (Culbreath et al., 2003) [15] . Despite their ecological and economic importance, knowledge of the diversity of thrips species across different legume host plants, their geographical expansion across tropical, subtropical, and temperate regions, their invasiveness, and their functional ecology remains incomplete. There is no single published work with a comprehensive synthesis. The present review aims to compile existing knowledge on species diversity, distribution, geographical patterns, ecology, economic importance, and destructive potential associated with pulse crops, and to provide a foundation for future integrated pest management interventions targeting these pests.

Taxonomy and classification of thrips 2.1 Taxonomic position and suborders

Thrips (Thysanoptera) are classified into two suborders, Terebrantia and Tubulifera. Major phytophagous thrips species belong to Terebrantia and are characterized by a saw-like ovipositor (terebra) in females, which helps them lay eggs inside plant tissues. In contrast, Tubulifera deposit eggs on the plant surface using a tube-like structure formed by the posterior end of the abdomen (Mound, 2005) [56] , and Phlaeothripidae is the only family in Tubulifera. More than 2000 species are present in Terebrantia, of which most of the agriculturally important species belong to the Family Thripidae (Mound and Morris, 2007) [55] . The key genera, such as Thrips, Frankliniella, Scirtothrips, Sericothrips and Megalurothrips, form pest complexes in pulse crops. The key thrips pest of legume crops is Megalurothrips usitatus worldwide (Mound and Marullo, 1996; Tyagi and Kumar, 2016) [54,88] . Many tubuliferan genera, such as Haplothrips, are phytophagous and some of which are mycophagous and predatory in nature (Mound and Marullo, 1996; Mound, 2005) [54,56] .

Cryptic species and species identification challenges

Thrips are minute in size, and slide mounting is essential to identify specific characteristics s of each genus (Mound and Kibby, 1998) [53,89] . The major taxonomic characters are longitudinal wing venation, antennal sensoria, ocellar setae, seta of pronotum, mesonotum and metanotum, abdominal tergite and ctenidia (Mound and Kibby, 1998; Moritz et al., 2004) [49,53,89] . Usually, the identification process requires expert knowledge and detailed analysis of slide-mounted specimens with suitable dichotomous keys. However, cryptic species within the same genus are challenging, and identifying them is cumbersome. Some characters show considerable variations, complicating reliable identification. Integration of morphological identification with molecular characterization can confirm these insects to the species level and is found to be more reliable (Hebert et al., 2003; Rebijith et al., 2014) [25,67] . The advent of modern approaches such as DNA barcoding using the cytochrome c oxidase subunit I (COI) gene, in silico sequence analysis, and the construction of phylogenetic trees using software such as MEGA has revolutionized thrips taxonomy (Marullo et al., 2020) [47] .

Computerized key-based photomicrographs and videos of thrips species are helpful to address morphological characters (Moritz et al., 2000) [48] . Semi-automated identification of biological objects can be achieved using artificial intelligence (Fedor et al., 2008) [19] . The morphometric analysis based on quantitative and qualitative characters, followed by principal component analysis, allows for distinguishing between cryptic species (Kucharczyk and Kucharczyk, 2009; Kucharczyk, 2010) [40,41] . Recently, complete genome sequencing has been performed for Frankliniella occidentalis and Thrips palmi (Rotenberg et al., 2020) [76] . These promising tools are reliable and easier for thrips identification than traditional procedures. Multilocus phylogenetics, high-throughput sequencing/metabarcoding, and integrative taxonomy are among the novel methods adopted for thrips identification (Batovska et al., 2021) [9] . The application of genomic and transcriptomic (omics) technologies was most recently utilized to understand thrips diversity. Long-read sequencing (PacBio/Illumina) and Hi-C scaffolding is employed to synthesize high-quality genomes for species such as Stenchaetothrips biformis (Hu et al., 2023) [26] .

Thrips species diversity in legumes 3.1 Thrips diversity worldwide

Earlier literature documents a diverse group of thrips species associated with pulse crops and approximately 60 thrips species are reported from legume production systems (Lewis, 1997; Mound and Marullo, 1996; Morse and Hoddle, 2006; Tyagi and Kumar, 2016) [44,45,54,88] . The thrips are attracted to legumes and infest leaves, flowers, and pods depending on growth stage, host plant cues, geographic patterns, altitude, and season. Tropical and subtropical regions support more abundant and diverse thrips species than temperate regions (Wang et al., 2014) [93] . Rainfall patterns also adversely affect thrips population buildup and the crop pest interactions are highly complex (Morsello et al., 2008) [51] . Major thrips species documented in legumes are listed in Table 1. Jandricic et al. (2024) [30] documented F. occidentalis and T. tabaci, as well as the newly invasive Thrips parvispinus, Frankliniella schultzei, and Echinothrips americanus, from an Ontario greenhouse during a survey from 2016 to 2024. Frankliniella occidentalis has been reported from at least sixty-nine countries (He et al. 2020; Fan et al. 2024) [17,23,24] . Thrips palmi is a key pest in legumes and a major tospovirus vector (WBNV and GBNV) on legumes in India. Distinct genetic signatures (microsatellite allele frequency distributions) were observed in T. palmi populations associated with legumes, indicating host-associated selection pressure on the species (Rehsawla et al., 2025) [68] . A multiyear study (2015-2018) on thrips associated with soybean highlighted the diverse assemblage of 20 thrips species across Eastern Europe, with two dominant species, viz., T. tabaci, Aeolothrips intermedius, and Thrips fuscipes. However, Kakothrips robustus, which specifically attacks crops in the Fabaceae, was absent in soybean (Twardowski et al., 2024) [86] . Molecular studies revealed the presence of cryptic species within Megalurothrips usitatus, T. palmi, and T. tabaci in legume cropping systems (Rebijith et al., 2014; Nault et al., 2014; Kadirvel et al., 2013) [31,59,67] . A diverse thrips fauna occurs in legumes globally, according to earlier literature.

Geographical distribution patterns 4.1 Tropical and subtropical regions

Sub-Saharan Africa, South Asia, and Southeast Asia are the major thrips diversity hubs, with the highest species richness and abundance. Tropical and subtropical ecosystems support several thrips species in legumes (Srinivasan et al., 2017) [83] . In Africa, Megalurothrips sjostedti is the major thrips species occurring in cowpea (Togola et al., 2019; Ngakou et al., 2008) [60,85] . Frankliniella schultzei was recorded as the most abundant and widely distributed species in Brazil, Colombia, Central America. Some invasive species, such as T. parvispinus and T. palmi, were reported tropical and subtropical regions than in temperate regions. Heliothrips ceylonicus, Frankliniella intonsa, Ceratothripoides claratris, Microcephalothrips abdominalis, Megalurothrips typicus and Craspedothrips minor were reported as new species occurring in vegetables and pulses in Central India (Akanand et al., 2026) [5] . Scirtothrips dorsalis, F. schultzei, T. palmi and Caliothrips indicus are the most common thrips species in groundnut (Kandakoor et al., 2014) [32] . The tropical and sub-tropical regions are favourable and support exceptionally high thrips diversity. These are major centers of diversity for several thrips genera (Mound, 2005) [56] . Dry weather (30.3 mm rainfall) with moderately high temperatures (15.6-28.2 °C) was found favourable to T. tabaci, whereas high humidity and rainfall negatively affected its population (Waiganjo et al., 2008) [92] . Most of the thrips species complexes recorded in tropical and subtropical legume systems are economically important and require timely interventions and management strategies.

Temperate regions

Species richness is comparatively low in legume crops across Europe, North America, and Temperate Asia. A severe winter with temperatures near 10 ºC did not support thrips populations in the temperate region, whereas high temperatures in June (25.8 ºC) led to thrips population peaks (Lopez et al., 2007) [99] . Neohydatothrips variabilis is the major thrips species infesting soybean in North America, which transmits Soybean Vein Necrosis Virus (SVNV) (Keough et al., 2016) [35] . Frankliniella occidentalis is another species that is invasive, causing significant crop losses in beans and peas across Europe and parts of Asia. In western and northern Europe, Kakothrips pisivorus has been documented as a host-specific thrips of pea and faba bean, whereas T. tabaci has been reported as a common and widely distributed species in parts of the Mediterranean region (Reitz, 2009) [71] . [67] A comprehensive barcoding survey of legumes, vegetables, and ornamentals in Bangladesh revealed that T. tabaci, F. schultzei, T. palmi, and C. indicus were confirmed using COI barcoding, and that T. parvispinus was recorded as invasive and newly documented in Bangladesh (Khatun et al., 2024) [36] . Thrips parvispinus was recorded in Florida and other warm US states and studies were conducted to manage this pest using conventional and biorational insecticides (Ataide et al., 2024) [8] . Thrips parvispinus was first recorded on pepper, and its detection is significant for legume production in Central America, as pepper and common bean are cultivated alongside in the same agroecological zones (Rodríguez Arrieta et al., 2025) [74] . It is now recorded across multiple continents globally distributed in legumes and other crops (Tyagi et al., 2015; Sridhar et al., 2021) [82,87] . A global invasion of T. parvispinus witnessed during the past few years urged tracing the invasion roots and genetic basis of the species' explosive global spread. The haplotype pattern revealed that all populations in Europe, the Americas, and Africa were derived from a single dominant haplotype, confirming a homogeneous founder effect (Ahmed et al., 2024) [4] . Stringent quarantine measures and surveillance systems are essential for detecting and managing invasive thrips species in legume agroecosystems. Jackai and Adalla, 1997) [10,28,72,78] . [7,23,24,70,95] . Thrips population peaks synchronized with the flowering stage (Twardowski et al., 2024) [86] . Jackai and Daoust (1986) [29] highlighted that thrips cause bronze discolouration and distortion on pods unfit for market.

Functional ecology of thrips in legume systems

Seasonal population dynamics

Abiotic factors, such as rainfall, temperature, and planting dates, strongly influence thrips population dynamics in legumes. An increase in thrips population was observed during the dry season, with low humidity, in tropical regions (Waiganjo et al., 2008) [92] . Temperate regions have peak populations during summer, coinciding with the optimal temperature during the flowering period (Lopez et al., 2007) [99] .

Thrips flower interactions

Complex ecological interactions are involved between legumes and flower-infesting thrips. Many thrips feed on pollen and floral exudates and hide inside papilionaceous flowers, which protect them from natural enemies and pesticide sprays. Beyond simple herbivory, they facilitate pollen exchange during foraging, thereby establishing a facultative mutualism. However, the negative effects of feeding on sap from flowers cause discolouration, flower deformation, and eventually flower bud abortion, which directly affect productivity by reducing pod set. The microhabitat within the closed floral structure is a potential barrier to effective control, as spray penetration into the flowers is comparatively low (Jackai and Daoust, 1986) [29] .

Thrips-virus interactions in legume crops 6.1 Orthotospoviruses transmitted by thrips in legumes

Thrips are vectors of orthotospovirus (Family: Tospoviridae). Orthotospoviruses such as tomato spotted wilt orthotospovirus (TSWV), groundnut ringspot orthotospovirus (GRSV), iris yellow spot orthotospovirus (IYSV) and watermelon silver mottle orthotospovirus (WSMoV) are enveloped, negative ssRNA viruses transmitted in a persistent propagative manner solely by 1 st instar larvae of thrips belonging to the family Thripidae. Twenty-nine tospoviruses are transmitted by sixteen thrips species globally (Ghosh et al., 2017) [20] . Tomato spotted wilt orthotospovirus (TSWV) affects a wide range of legumes and is transmitted by F. occidentalis (Rotenberg et al., 2015) [77] , whereas IYSV is transmitted by T. tabaci (Pandi et al., 2024) [63] . Cowpea, groundnut, and soybean were found affected by WSMoV transmitted by T. palmi (Mou et al., 2021; Pradeep et al., 2024) [52,66] , whereas Groundnut bud necrosis orthotospovirus (GBNV) transmits mungbean, cowpea, french bean, field pea, urdbean and wild Vigna species are severely affected by (Akram et al., 2026) [6] . Groundnut Ringspot Virus (GRSV) is transmitted by F. schultzei (Riley et al., 2011; Pappu et al., 2009) [64,73] . The major virus in soybean is Soybean Vein Necrosis Virus (SVNV), which is transmitted by Neohydatothrips variabilis (Rotenberg et al., 2015; Schwartz et al., 2020) [77,100] . Major thrips species that transmit viruses associated with legumes cause substantial yield losses and require integrated pest management strategies to reduce thrips populations below the economic threshold.

Transmission efficiency of the thrips species

The epidemiology of orthotospovirus diseases is directly influenced by thrips abundance in legumes and susceptibility of plants (Pappu et al., 2009; Rotenberg et al., 2015) [64,77] . Young seedlings are affected severely compared to older plants. A high thrips population is directly linked to rapid virus infection (Riley et al., 2011) [73] . Furthermore, environmental factors can influence virus transmissions and thrips population (Rotenberg et al., 2015; Keough et al., 2018) [35,77] . Resistance varieties, cultural control, and insecticide applications are integrated to effectively prevent virus transmission through vectors.

Damage mechanisms and economic impact 7.1 Direct feeding damage

Asymmetric rasping and sucking mouth parts with reduced mandibles leave lacerations on plant tissue caused by rupture of epidermal and mesophyll cells. The silvery streaks and bronzing, combined with leaf distortion and malformation, cause defoliation and stunted growth in young plants (Kumar and Omkar, 2021) [42,43] . Flowers are significantly affected by thrips feeding, with discolouration and malformation ultimately leading to abortion of flower buds and crop failure.

Yield loss assessment

A possible yield loss from 80-100% was reported from previous studies (Singh and Allen, 1980; Nault et al., 2006) [81, 101] . The simultaneous occurrence of thrips complexes in crop fields, their detection, identification, direct and indirect damage assessment, resistance development, and host plant resistance are attributed to the complexity of thrips management in legumes (Riley et al., 2011) [73] . EIL was established at seven thrips per inflorescence for M. sjostedti infesting cowpea (Nabirye et al., 2003) [58] . Similarly, a loss of 36.65% of pods was experienced on green beans due to thrips damage (Duff et al., 2014) [16] . [12,69,79] . Field sanitation is adopted to eliminate potential breeding sites of thrips. A chemotactic approach based on a push-pull strategy using an alarm pheromone and an aggregation pheromone was followed against western flower thrips (Kim et al., 2023) [37] .

Drip irrigation reduces thrips populations compared to sprinkler irrigation. The cultural practices minimise chemical interventions in integrated pest management (Rodriguez et al., 2023) [75] . Optimising these control tactics based on specific crop and environmental factors is essential for effective thrips management.

Host plant resistance

Breeding and use of resistant varieties are widely considered sustainable approaches for managing thrips in legumes. It is the most economically accessible management strategy for marginal farmers. There are three resistance mechanisms associated with legumes to sustain yields despite thrips infestation, viz., antixenosis, antibiosis, and tolerance (Painter, 1951) [62] . [11,57] .

Biological control

Predatory mites (Amblyseius cucumeris, Neoseiulus barkeri and Typhlodromips montdorensis) and predatory thrips, Orius sp., Geocoris sp., entomopathogenic fungi (Beauveria bassiana, Metarhizium anisopliae and Lecanicillium lecanii) and entomopathogenic nematodes are natural enemies documented against thrips. Beauveria bassiana, predatory mites and pirate bugs are important bioagents against thrips infesting legumes (Chi et al., 2024) [14] . Biological control is considered variable, depending on environmental factors. Reduced pesticide use, timing of applications, and maintenance of landscape heterogeneity can conserve natural enemies.

Chemical control and insecticide resistance

The most important management tool for thrips in the legume production system is synthetic insecticides. Neonicotinoids, organophosphates, and pyrethroids are among the insecticide groups used against thrips. Seed treatment with neonicotinoids, such as thiamethoxam, is widely practised. Spinetoram and spinosad are reported as the most effective insecticides for M. usitatus. Moreover, broflanilide, emamectin benzoate and cyantraniliprole were also recorded as alternative insecticides against this species (Peng et al., 2025) [65] . Seed treatment with thiamethoxam and imidacloprid was found effective against M. sjostedti infesting cowpea (Kaushik et al, 2015) [3,4] . However, insecticide resistance is a major threat in thrips management. Even new insecticide molecules are reported to be less effective against thrips populations. Field evolved resistance of bean flower thrips, M. usitatus, to broflanilide with LC 50 values ranging from 7804.77 to 20 594.08 mg/L (Kang et al., 2026) [33] . Resistant management strategies, including insecticide rotation with insecticides of different modes of action, leaving refugia for susceptible populations, and integrating with non-chemical methods, are essential to slow down resistance development.

Conclusion

Substantial knowledge gaps remain in the basic understanding of thrips diversity and management, and these need to be addressed. Systematic documentation of thrips diversity in understudied legume cultivating regions, identification of cryptic species complexes within genera by molecular characterisation using sophisticated techniques, elucidation of underlying crop-pest interaction mechanisms, detailed epidemiological studies of virus diseases, investigation of genetic basis of host plant resistance, assessment of the effect of climatic factors on thrips population dynamics and evaluation of resistance development in legume thrips are identified as important future line of research. Thrips are a major pest associated with legumes worldwide. There are diverse thrips species distributed in tropical, subtropical and temperate regions. Most thrips are highly polyphagous. These pests are responsible for significant economic damage and yield losses through direct feeding damage and indirect damage through virus transmission keeps them the most challenging pests of legumes. Species identification, using both morphological and molecular methods, is integrated to confirm cryptic and major species, which is essential for decision-making and effective management.

An integrated management approach is required to achieve effective, environmentally safe population reduction below ETL. It includes host plant resistance, cultural control, biological control and chemical management. Breeding of resistant varieties using modern genomic tools and multiple field evaluations of varietal performance against thrips accelerates their development and deployment. The environmental factors influencing thrips population and community dynamics in legume ecosystems, and the effects of natural enemies, determine thrips buildup in legume ecosystems.

The global threat posed by invasive species like T. parvispinus and T. palmi, and their expansion, urges immediate action to reduce thrips infestations in legume systems. Rising concerns about the devastating effects of the thrips species on yield losses and crop productivity demand effective pest management strategies. This review provides a foundation of baseline data on thrips diversity in legumes over the last ten years and helps to catalyse further research on sustainable thrips management to safeguard legume production and food security

Acknowledgement

AcknowledgementsThe authors thank the Department of Agricultural Entomology, College of Agriculture, Vellanikkara

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