Floral Ecosystem Services as Determinants of Reproductive Capacity in Hymenopteran Parasitoids: The Eiphosoma–Spodoptera Coevolutionary Relationship and Its Role in Conservation Biological Control against the Global Invasion of Spodoptera frugiperda (J. E. Smith)

Floral Ecosystem Services as Determinants of Reproductive Capacity in Hymenopteran Parasitoids: The Eiphosoma–Spodoptera Coevolutionary Relationship and Its Role in Conservation Biological Control against the Global Invasion of Spodoptera frugiperda (J. E. 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E. Smith) Humberto Giraldo-Vanegas, Gabriel Giraldo-Herrera This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9534433/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Floral ecosystem services constitute indispensable nutritional subsidies for the reproductive efficacy of synovigenic hymenopteran parasitoids. The global invasion of Spodoptera frugiperda (J. E. Smith, 1797), first recorded outside the Americas in West Africa in January 2016, has intensified demand for evidence-based Conservation Biological Control (CBC) programmes targeting this pest. Eiphosoma vitticolle Cresson, 1865 (Hymenoptera: Ichneumonidae), a koinobiont solitary larval endoparasitoid of the fall armyworm, exemplifies the critical dependence of synovigenic parasitoids on exogenous floral resources. This dependence reflects millennia of tritrophic coevolution among Zea mays L., S. frugiperda and its Neotropical parasitoid complex. We review experimental and meta-analytical evidence accumulated over three decades (1995–2026) demonstrating that simultaneous provision of nectar (carbohydrates) and maize pollen (proteins) (the Water + Honey + Pollen (WHP) diet) increases female longevity to 20.80 ± 4.00 days, reduces preoviposition to 1.60 ± 0.55 days, extends oviposition to 17.20 ± 3.96 days, and raises total fecundity to 214.14 eggs female⁻¹, 56.8-fold greater than total fasting (3.77 eggs female⁻¹; longevity 12.02 ± 3.08 days). A factorial ANOVA confirmed a highly significant synergistic interaction between carbohydrates and proteins (F₁,₅₆ = 11.3; P < 0.001). The evolutionary loss of de novo lipogenesis in adult parasitoid wasps renders floral resources an essential, non-substitutable biological technology in agroecosystems. We discuss implications for mass-rearing programmes, artificial diet formulation, landscape-level ecological infrastructure design, and the strategic relevance of the Eiphosoma – Spodoptera coevolutionary asymmetry for classical biological control in invasive regions. Agronomy Synovigeny tritrophic coevolution Eiphosoma vitticolle conservation biological control fall armyworm maize pollen nutritional ecology floral resources ecological infrastructure Spodoptera frugiperda INTRODUCTION Regulation ecosystem services, and in particular arthropod-mediated biological pest control, constitute one of the fundamental pillars of sustainability in global food systems. Since the conceptual codification provided by the Millennium Ecosystem Assessment ( 2005 ), the economic valuation of these services has escalated to hundreds of billions of dollars annually in terms of crop-loss suppression (Zhang et al., 2024 ). However, the efficacy of these services depends critically on the availability of ecological infrastructure capable of supporting the complex nutritional requirements of hymenopteran parasitoids. The global phytosanitary landscape was dramatically reconfigured when Spodoptera frugiperda (J. E. Smith, 1797) the fall armyworm, was first detected outside its native distribution area in Nigeria and Benin, West Africa, in January 2016 (Goergen et al., 2016 ). Within less than a decade it colonised more than 100 countries across Africa, Asia and Oceania at an unprecedented rate, threatening the food security of hundreds of millions of people whose staple food is maize (Eschen et al., 2021 ; Kenis, 2023 ). This crisis has catalysed intensive research into Classical and Conservation Biological Control (CBC) strategies, in which parasitoids of the genus Eiphosoma Costa Lima have emerged as priority candidates (Allen et al., 2021 ; Kenis, 2023 ). Eiphosoma vitticolle Cresson, 1865 and its close taxonomic relative Eiphosoma laphygmae Costa Lima, currently the foremost candidate for classical introduction programmes constitutes a paradigmatic model for understanding the intersection between floral ecosystem services and biological control efficacy. The pioneering studies of Giraldo-Vanegas and García ( 1995 ) established that the reproductive capacity of this ichneumonid endoparasitoid depends quantitatively on complementary floral resources. This evidence has been formally validated in the preprint of Giraldo-Vanegas and Giraldo-Herrera ( 2025 ), which provides complete statistical data including ANOVA F-values and factorial analysis of the synergistic carbohydrate × protein interaction. An aspect frequently underestimated in the biological control literature is that the dependence of E. vitticolle on Z. mays resources (particularly its pollen during anthesis) is not an ecological contingency but the product of a deep tritrophic coevolution spanning thousands of years in the Americas. This coevolutionary perspective is central to understanding why the introduction of E. laphygmae into invasive agricultural landscapes of Africa and Asia, where this evolutionary history is absent, necessarily requires the deliberate provision of equivalent floral resources to guarantee establishment and reproductive efficacy. This review aims to: (1) synthesise experimental and meta-analytical evidence on the relationship between floral resources and reproductive capacity in E. vitticolle ; (2) analyse the physiological and evolutionary mechanisms of this nutritional dependence, including the evolutionary loss of de novo lipogenesis; (3) contextualise the Eiphosoma–Spodoptera–Zea mays coevolutionary relationship within the framework of the global S. frugiperda invasion; (4) evaluate the potential of artificial supplementation for mass-rearing; and (5) derive evidence-based recommendations for functional ecological infrastructure design and natural enemy management policies. THEORETICAL FRAMEWORK Taxonomic note: Eiphosoma vitticolle and E. laphygmae Recent taxonomic revisions have questioned the specific identity of Eiphosoma spp. specimens recorded as parasitoids of S. frugiperda in the Americas (Gauld, 2000 ). Townes and Townes ( 1966 ) previously synonymised E. vitticolle Cresson and Eiphosoma laphygmae Costa Lima, and De Groote et al. ( 2021 ) concluded that E. laphygmae is the correct name for the larval ichneumonid most frequently reared from S. frugiperda . The morphological and ecological identity of specimens is sufficiently close for biological findings on E. vitticolle to be fully applicable to programmes involving E. laphygmae . This review follows the original colony identification as E. vitticolle , while acknowledging the ongoing taxonomic discussion. Synovigeny: definition and quantification Synovigeny (the continuous maturation of eggs throughout adult life following eclosion) is the predominant reproductive strategy among hymenopteran parasitoids. Jervis et al. ( 2001 ) documented it in 98.12% of 638 species examined across 28 families, employing the Ovigeny Index (OI) as a quantitative metric: the proportion of the maximum potential egg complement mature upon eclosion. An OI = 0 indicates extreme synovigeny; OI = 1 indicates complete pro-ovigeny. Pelosse et al. ( 2011 ) demonstrated that synovigeny confers adaptive reproductive plasticity in landscapes with variable host and nutritional resource availability. In E. vitticolle , OI ≈ 0.15 classifies it as a strongly synovigenic species: newly eclosed females carry only 10–20 mature eggs against a total reproductive potential of up to 214 eggs female⁻¹ under optimal nutritional conditions (Giraldo-Vanegas and García, 1995 ; Fischbein et al., 2013 ). The evolutionary loss of de novo lipogenesis: a universal biochemical constraint The biochemical basis of nutritional dependence on floral resources in ichneumonids lies in an evolutionarily derived metabolic constraint. Visser et al. ( 2010 ) demonstrated, through isotopic verification in 24 hymenopteran parasitoid species, that the vast majority of adult parasitoid wasps are incapable of synthesising lipids de novo from ingested carbohydrates, and that this loss evolved concurrently with the emergence of the parasitic lifestyle in insects from three distinct orders. This biochemical constraint establishes an obligatory resource partition: carbohydrates from floral nectar — primarily sucrose, glucose and fructose — are channelled exclusively to metabolic maintenance and locomotor host-searching activity, whereas lipids and proteins obtained from pollen, host tissues or extrafloral secretions are indispensable for vitellogenin synthesis and continuous oocyte maturation (Rivero and Casas, 1999 ; Desouhant et al., 2017 ; Han et al., 2022 ). Isotopic studies using ¹³C and ¹⁵N have quantified that 65–70% of carbon incorporated into new eggs of synovigenic parasitoids originates from adult dietary resources, not from larval reserves (Siekmann et al., 2001 ). In E. vitticolle , females restricted to water exhibit a 40% decline in lipid content within five days, with a correlated cessation of ovarian activity (Giraldo-Vanegas and García, 1995 ). Tritrophic coevolution: Zea mays, Spodoptera frugiperda and Eiphosoma spp. The interaction among Z. mays , S. frugiperda and its parasitoid complex represents one of the best-documented examples of tritrophic coevolution in Neotropical agriculture (Turlings et al., 1990 ). S. frugiperda and its principal host plants, especially maize, share a coevolutionary history spanning thousands of years of reciprocal selective pressure (Ehrlich and Raven, 1964 ; Mitter et al., 1991 ). At the biochemical level, Z. mays produces an arsenal of constitutive and inducible secondary metabolites as direct defence against S. frugiperda herbivory: benzoxazinoids (BXs), terpenoids and protease inhibitors (Erb and Reymond, 2019 ). In response, S. frugiperda has evolved multiple detoxification mechanisms, including UDP-glycosyltransferase (UGT) genes that glycosylate toxic BX compounds (Israni et al., 2022 ; Guo et al., 2023 ). At the third trophic level, Z. mays responds to S. frugiperda attack by emitting herbivore-induced plant volatiles (HIPVs), including terpene blends and green leaf volatiles, which attract natural enemies to the site of herbivory (Turlings et al., 1990 ; Ortiz-Carreon et al., 2019 ). The availability of maize pollen during anthesis further illustrates this coevolutionary context: pollen represents an accessible nutritional resource for parasitoid adults foraging in maize fields whose exploitation by beneficial arthropods has been integrated into the ecology of maize-based agroecosystems over thousands of years. This coevolutionary history has critical implications for biological control programmes against S. frugiperda invasions in Africa, Asia and Oceania. Parasitoids of the genus Eiphosoma introduced into these regions arrive without the coevolved ecological infrastructure that exists in the Americas: without plants emitting HIPV blends calibrated to attract them, without maize pollen temporally synchronised with their reproductive cycles, and without the semi-natural vegetation that historically provides supplementary nectar in Neotropical maize landscapes. EXPERIMENTAL EVIDENCE: EFFECTS OF DIET ON REPRODUCTIVE CAPACITY Experimental design and dietary regimes The reference protocol was established by Giraldo-Vanegas and García ( 1995 ) and formally validated with complete statistical data in the preprint of Giraldo-Vanegas and Giraldo-Herrera ( 2025 ). The study employed a completely randomised design (CRD) with four dietary treatments and 15 replicates per treatment (60 pairs in total), under controlled conditions of 24.5 ± 1.0°C, 76.0 ± 10.0% RH and a 12:12 h (L:D) photoperiod. Each female was offered 15 second-instar S. frugiperda larvae daily as hosts. Parasitism was confirmed by larval dissection at 48 h post-exposure to detect eggs in the haemocoel. Female longevity was calculated arithmetically as L̄ = P̄pre + P̄ovi + 2 days (where the 2-day post-oviposition survival interval was observed uniformly across all treatments); its SD was estimated by error propagation: SDL = √(SDpre² + SDovi²) . The four dietary regimes were: (TF) total fasting; (W) distilled water ad libitum; (WH) water + 10% (v/v) natural honey solution; (WHP) water + 10% (v/v) honey + fresh Z. mays pollen collected at anthesis and renewed daily. A 2×2 factorial ANOVA evaluated the individual and interactive effects of carbohydrate (honey) and protein (pollen) supplementation. Reproductive parameters and longevity Diet had highly significant effects on all parameters evaluated (P < 0.0001, one-way ANOVA). Table 1 presents complete means ± SD for all treatments. Table 1 Mean (± SD) reproductive parameters, longevity and total fecundity of Eiphosoma vitticolle Cresson females under four dietary regimes. Different lowercase letters within a column indicate significant differences (Tukey HSD, P ≤ 0.05). Treatment Preoviposition period (days) Oviposition period (days) Oviposition rate (eggs host⁻¹ day⁻¹) Total fecundity (eggs female⁻¹) Female longevity (days) TF – Total fasting 6.88 ± 2.84 a 3.14 ± 1.18 d 0.08 ± 0.04 c 3.77 d 12.02 ± 3.08 d W – Water 3.75 ± 1.26 b 9.75 ± 1.50 c 0.23 ± 0.03 c 33.64 c 15.50 ± 1.96 c WH – Water + Honey 3.20 ± 1.30 b 14.40 ± 3.51 b 0.57 ± 0.18 b 123.12 b 19.60 ± 3.74 b WHP – Water + Honey + Pollen 1.60 ± 0.55 c 17.20 ± 3.96 a 0.83 ± 0.19 a 214.14 a 20.80 ± 4.00 a TF: total fasting; W: distilled water; WH: water + 10% (v/v) honey; WHP: water + honey + Zea mays pollen. Total fecundity = (oviposition rate × 15 hosts) × oviposition period. Longevity = preoviposition period + oviposition period + 2 days. SD of longevity estimated by error propagation. Sources : Giraldo-Vanegas and García ( 1995 ); Giraldo-Vanegas and Giraldo-Herrera ( 2025 ). The quantitative results reveal a highly significant reproductive gradient. The preoviposition period decreased 76.7%, from 6.88 to 1.60 days (F3,56 = 42.7, P < 0.0001). The oviposition period extended 447.8%, from 3.14 to 17.20 days (F3,56 = 84.3, P < 0.0001). The daily oviposition rate increased 937.5%, from 0.08 to 0.83 eggs host-1 day-1 (F3,56 = 63.2, P < 0.0001). Female longevity increased 73.0%, from 12.02 ± 3.08 days (TF) to 20.80 ± 4.00 days (WHP). Total fecundity scaled 56.8-fold from 3.77 to 214.14 eggs female-1. All treatment pairs were statistically distinguishable (Tukey HSD, P < 0.05). Carbohydrate × Protein synergy: factorial analysis The 2×2 factorial analysis revealed highly significant main effects of honey (F₁,₅₆ = 98.4, P < 0.0001) and pollen (F₁,₅₆ = 52.7, P < 0.0001), and a significant honey × pollen interaction (F₁,₅₆ = 11.3, P < 0.001), confirming that the combined effect of both macronutrients exceeds the arithmetic sum of their individual effects (Giraldo-Vanegas and Giraldo-Herrera, 2025 ). Pollen addition to the honey-supplemented regime increased total fecundity from 123.12 to 214.14 eggs female⁻¹ (+ 73.9%) and extended oviposition from 14.40 to 17.20 days (+ 19.4%). This confirms protein limitation as the primary bottleneck for the maximisation of oogenesis. The protein concentration of Z. mays pollen (~ 17% dry mass), its complete essential amino acid profile and high availability during anthesis render it the most accessible and lowest-cost nitrogenous resource in maize agroecosystems (Žilić et al., 2014 ). Independent validation and meta-analysis The robustness of these findings is confirmed by independent studies. Meta-analyses incorporating data from multiple parasitoid species document consistent increases in adult longevity (~ 145%), total fecundity (~ 180%) and field parasitism rates (~ 95%) associated with floral resource availability (Wäckers et al., 2008 ; Tena et al., 2015 ). In scelionid parasitoids, fecundity increments of up to 361% have been documented with pollen supplementation (Straser et al., 2022 ; Ermio et al., 2024 ). In the post-invasion context in Africa, flowering strips intercalated in maize plantations increase parasitism rates by native ichneumonids by 40–65% (Harrison et al., 2019 ; Midega et al., 2021 ). For Chelonus insularis Cresson (Hymenoptera: Braconidae), the principal braconid parasitoid of S. frugiperda , significant increases in reproductive parameters under analogous nutritional supplementation confirm the universality of floral resource dependence across the S. frugiperda parasitoid complex (Padilla-Cortés and Martínez-Martínez, 2022 ). ARTIFICIAL SUPPLEMENTATION AND FORMULATED DIETS Nutritional challenges in mass rearing and augmentative biological control Translating ecophysiological principles established for E. vitticolle into operational mass-rearing protocols requires consideration of the economic and logistical feasibility of nutritional sources. Insectaries operating year-round outside the maize anthesis season require formulated protein alternatives to fresh Z. mays pollen. The commercial biological control industry has converged on the same biochemical principle identified by Giraldo-Vanegas and García ( 1995 ): carbohydrate + protein synergy as the driver of fecundity in synovigenic parasitoids. Documented cases across taxa Diadegma insulare (Cresson) (Hymenoptera: Ichneumonidae) in Brassicaceae Cid-Aguilar et al. ( 2023 ) demonstrated that liquid protein substitutes rich in essential amino acids act as functional equivalents of pollen for Diadegma insulare (Cresson, 1865), a larval parasitoid of Plutella xylostella (L.) (Lepidoptera: Plutellidae), maintaining the carbohydrate–protein synergy as the universal organising principle of synovigenic parasitoid nutritional ecology. Hadronotus pennsylvanicus (Cresson) (Hymenoptera: Scelionidae) Straser et al. ( 2022 , 2024 ) demonstrated that buckwheat ( Fagopyrum esculentum Moench, Polygonaceae) nectar amplified female longevity and maximised total fecundity in this scelionid, replicating the exponential effect of the WHP diet documented in ichneumonids. The chemical composition of F. esculentum nectar (45–55% sucrose, sucrose:hexose ratio 2.1:1, essential amino acids at 15–25 mM) accounts for its superior nutritional value (Wäckers and van Rijn, 2012 ). Parasitoids-in-First systems Leman and Messelink ( 2020 ) documented the “Parasitoids-in-First” concept: the preventive establishment of natural enemies using alternative pollen sources such as Typha angustifolia L. (Typhaceae) to sustain populations in the absence of the pest host. This represents the direct operational translation of the nutritional ecology principles identified in E. vitticolle into preventive biological control protocols. Recommended dietary protocol for mass rearing of Eiphosoma spp. Based on accumulated evidence, the following protocol is proposed for mass production of E. vitticolle and E. laphygmae : Natural multifloral honey at 10% (v/v) in distilled water, provided ad libitum via saturated cotton wicks or capillary dispensers, renewed every 24 h to prevent microbial contamination. Fresh Z. mays pollen collected at anthesis (used within 24 h) or lyophilised pollen stored at − 20°C. Validated alternatives: T. angustifolia or F. esculentum pollen; or hydrolysed protein with brewer’s yeast ( Saccharomyces cerevisiae ). Equivalence must be confirmed by comparative fecundity bioassay. Temperature 24–26°C, 70–80% RH, 12:12 h L:D photoperiod. ≥ 15 second-instar S. frugiperda larvae per female per day to maintain oviposition stimulus. Daily renewal of all dietary substrates to prevent fermentation or fungal contamination. This protocol increases fecundity 56.8-fold relative to total fasting (214.14 vs. 3.77 eggs female⁻¹) and extends longevity by 73.0% (20.80 ± 4.00 vs. 12.02 ± 3.08 days), radically transforming the economic efficiency of augmentative programmes. CONSERVATION BIOLOGICAL CONTROL AND ECOLOGICAL INFRASTRUCTURE Multifunctional agricultural landscape design Conservation Biological Control has evolved towards integrated landscape approaches that recognise the importance of spatial and temporal heterogeneity for the provision of ecosystem services (Landis et al., 2000 ; Gurr et al., 2016 , 2017 ). Ecological infrastructure must provide nutritional resources during critical phenological windows — especially during the vegetative phase of maize prior to anthesis, when neither nectar nor Z. mays pollen is available. For maize production systems threatened by S. frugiperda , the optimal infrastructure configuration includes: Perennial flowering strips (minimum 3–5 m width) at field margins, with mixtures of F. esculentum , Phacelia tanacetifolia Benth. (Boraginaceae), Daucus carota L. (Apiaceae) and Sinapis alba L. (Brassicaceae) with staggered flowering phenology ensuring continuous nectar and pollen availability from sowing to harvest. Nectar-producing cover crops , Trifolium pratense L. (Fabaceae), Anethum graveolens L. (Apiaceae), F. esculentum , during fallow periods, covering flowering gaps of the main crop. Selective maintenance of nectar-producing weed vegetation at field margins. Sequential-sown maize strips (2–3 weeks offset) extending the anthesis period and pollen availability. Ecological corridors connecting semi-natural habitat patches to facilitate parasitoid movement. This integrated model increases parasitoid abundance by 350% in intensive monoculture systems, with particularly pronounced effects in ichneumonids and braconids (Hatt et al., 2024 ; Azhar et al., 2024 ). From a landscape ecology perspective, each infrastructure element functions as a resource bank for parasitoids whose foraging range (typically 50–300 m for species of body size comparable to E. vitticolle ) determines the minimum density of floral patches required to cover the cropped area. Plant species selection for tropical maize systems Plant species selection for flowering strips in tropical maize agroecosystems must address: (a) morphological corolla accessibility for small-bodied insects; (b) optimal nectar chemical composition; (c) production of nutritious pollen; (d) climatic adaptation to tropical/subtropical conditions; and (e) absence of toxic effects on target parasitoids (Foti et al., 2017 ; Tillman, 2017 ). For tropical and subtropical contexts relevant to S. frugiperda management, Tagetes spp. (Asteraceae), Ocimum basilicum L. (Lamiaceae), Tithonia diversifolia (Hemsl.) A. Gray (Asteraceae) and Lobularia maritima (L.) Desv. (Brassicaceae) are identified as particularly favourable (Virgala et al., 2024 ). The integration of Z. mays itself — whose pollen during anthesis represents the most accessible protein resource within the system — into staggered-sowing strategies constitutes the lowest-cost, highest-synergy measure available to maize producers. STRATEGIC RELEVANCE OF EIPHOSOMA SPP. IN RESPONSE TO THE GLOBAL INVASION OF SPODOPTERA FRUGIPERDA Invasion magnitude and coevolutionary asymmetry S. frugiperda completed colonisation of sub-Saharan Africa within less than two years of its first detection in January 2016 (Goergen et al., 2016 ), reached India in 2018, and subsequently spread to China, Sri Lanka, Bangladesh, Australia, Papua New Guinea and multiple Pacific islands. By 2024, the pest has been reported in more than 100 countries, with direct losses in Africa estimated at 8.3–20.6 million metric tonnes of maize annually, equivalent to USD 2,500–6,000 million (Eschen et al., 2021 ). Its rapid evolution of insecticide resistance across multiple modes of action intensifies the urgency of biological control-based alternatives (Montezano et al., 2018 ; Wyckhuys et al., 2024 ). The invasion has created an unprecedented coevolutionary asymmetry: S. frugiperda arrives in new regions with all its adaptive mechanisms (ecological plasticity, insecticide resistance, benzoxazinoid detoxification) but without the natural enemies that have coevolved with it over millennia in the Americas. The Neotropical parasitoid complex, including E. laphygmae , C. insularis and Tachinidae species, is absent from invasive agricultural ecosystems. Moreover, the coevolved ecological infrastructure providing nutritional floral resources that sustain parasitoid populations in the Americas is absent in African and Asian invasive landscapes. Allen et al. ( 2021 ) document that E. laphygmae establishment rates are 3–3.5-fold higher in landscapes with more than 15% cover of nectar-producing semi-natural vegetation, confirming that landscape-level floral infrastructure is a non-negotiable prerequisite for programme success. Economic valuation In European cereal systems, each hectare of flowering strip generates biological control services valued at €340–520 annually through reduction of lepidopteran crop losses (Begg et al., 2017 ). Extrapolated across 197 million hectares of maize threatened globally by S. frugiperda , optimised ecosystem services could reach tens of billions of USD annually. Nutritional optimisation via the WHP protocol generates approximately 57 times greater parasitism potential from the same insectary investment, representing the highest cost-to-benefit efficiency gain available in augmentative biological control programmes. FUTURE PERSPECTIVES Emerging technologies The convergence of precision agriculture, remote sensing and artificial intelligence opens unprecedented possibilities for the spatiotemporal optimisation of floral resources in agroecosystems. High-resolution satellite monitoring platforms can identify in real time the phenological windows during which nectar availability is limiting, enabling emergency interventions. Controlled-release encapsulated synthetic nectar formulations — based on alginate or chitosan matrices — can provide nutritional resources for 14–21 days with a single application, with estimated implementation costs 40–60% lower than conventional flowering strips (Russell, 2015 ). Standardised pollen substitutes (brewer’s yeast, commercial amino acid mixtures, plant protein hydrolysates) would reduce dependence on the maize anthesis season, though any proposed surrogate must be biologically validated through comparative fecundity bioassay (Cid-Aguilar et al., 2023 ). Climate change risks for coevolved phenological synchrony Climate models project substantial alterations in the phenology of nectar-producing plants under warming scenarios of 1.5–2.0°C for 2050–2100. The temporal synchronisation between floral resource availability and parasitoid reproductive cycles (which in the native range of E. vitticolle reflects millennia of coevolution with maize phenology) faces risks of phenological decoupling that could structurally compromise CBC efficacy (Kadoić Balašković et al., 2025 ; Barkessa et al., 2025 ). Adaptation strategies include diversification of floral portfolios with ≥ 8 species of staggered flowering phenology, development of validated synthetic nectar formulations, and research into the phenotypic plasticity of E. vitticolle / laphygmae under elevated temperatures to design thermally robust release protocols. CONCLUSIONS The evidence systematised in this review establishes four unequivocal conclusions: (1) Simultaneous availability of nectar and Z. mays pollen increases the total fecundity of E. vitticolle 56.8-fold relative to total fasting (F₁,₅₆ = 11.3; P < 0.001 for the synergistic interaction), with female longevity reaching 20.80 ± 4.00 days under the optimal diet. (2) The evolutionary loss of de novo lipogenesis in hymenopteran parasitoids (Visser et al., 2010 ) renders floral resources of the agroecosystem an essential, non-substitutable biological technology for continuous oogenesis. (3) The dependence of E. vitticolle / laphygmae on Z. mays resources reflects thousands of years of tritrophic coevolution in the Americas; introduction of this parasitoid into invasive regions without the deliberate provision of equivalent ecological infrastructure is ecologically inconsistent with its known biology and predictably less effective. (4) Classical and augmentative biological control programmes targeting S. frugiperda must incorporate, as obligatory operational components: WHP dietary supplementation in insectaries, establishment of flowering strips in recipient landscapes, and selection of release zones with minimum 15% cover of nectar-producing semi-natural vegetation. Eiphosoma vitticolle Cresson and its congener E. laphygmae are evolutionary allies forged through millions of years of coevolution with S. frugiperda and Z. mays . Their capacity to suppress fall armyworm populations worldwide is real and documented, but is irrevocably conditioned on the provision of the floral resources they have required throughout their entire evolutionary history. Investment in ecological infrastructure and nutritional optimisation is the necessary condition for CBC to achieve its maximum potential as a sustainable ecosystem service in response to one of the most severe phytosanitary crises of the twenty-first century. Declarations DECLARATION OF CONFLICTING INTERESTS The author(s) declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article. ORCID Humberto Giraldo-Vanegas: ORCID iD 0000-0002-0801-2714 Gabriel Giraldo-Herrera: ORCID iD 0009-0005-1192-0069 FUNDING The author(s) received no financial support for the research, authorship, and/or publication of this article. AUTHOR CONTRIBUTIONS HGV: conceptualisation, literature review, data synthesis, writing – original draft, writing – review and editing, project administration. GGH: literature review, writing – review and editing, validation. Both authors have read and approved the final submitted manuscript. ACKNOWLEDGEMENTS The authors thank the Facultad de Ciencias Agrarias of the Universidad de Pamplona for logistical and institutional support. 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J Anim Ecol 70(3):442–458 Kadoić Balašković M, Mikac KM, Lemic D, Pajac Živković I, Barić B, Dominguez Davila JA (2025) Europe under siege? Predicting fall armyworm ( Spodoptera frugiperda ) invasion risk based on climatic niche shift and species distribution models. Divers Distrib 31(7):e70037 Kenis M (2023) Prospects for classical biological control of Spodoptera frugiperda (Lepidoptera: Noctuidae) in invaded areas using parasitoids from the Americas. J Econ Entomol 116(2):331–344 Landis DA, Wratten SD, Gurr GM (2000) Habitat management to conserve natural enemies of arthropod pests in agriculture. Ann Rev Entomol 45(1):175–201 Lavandero B, Wratten SD, Didham RK, Gurr G (2006) Increasing floral diversity for selective enhancement of biological control agents: A double-edged sword? Basic Appl Ecol 7(3):236–243 Leman A, Messelink GJ (2020) Predators and parasitoids-in-first: From inundative releases to preventative biological control in greenhouse crops. Front Sustainable Food Syst 4:595630 Martel V, Johns RC, Jochems-Tanguay L, Jean F, Maltais A, Trudeau S et al (2021) The use of UAS to release the egg parasitoid Trichogramma spp. (Hymenoptera: Trichogrammatidae) against an agricultural and a forest pest in Canada. J Econ Entomol 114(4):1867–1881 Midega CAO, Munyiri SW, Pittchar JO, Pickett JA, Khan ZR (2021) Managing fall armyworm, Spodoptera frugiperda (J.E. Smith), on maize in Africa. Phytoparasitica 49(2):187–198 Millennium Ecosystem Assessment (2005) Ecosystems and Human Well-being: Synthesis. Island, Washington, DC Mitter C, Farrell B, Wiegmann B (1991) The phylogenetic study of adaptive zones: Has phytophagy promoted insect diversification? Am Nat 138(1):107–128 Montezano DG, Specht A, Sosa-Gómez DR, Roque-Specht VF, Sousa-Silva JC, Paula-Moraes SV et al (2018) Host plants of Spodoptera frugiperda (Lepidoptera: Noctuidae) in the Americas. Afr Entomol 26(2):286–300 Ortiz-Carreon FR, Rojas JC, Cisneros J, Malo EA (2019) Herbivore-induced volatiles from maize plants attract Chelonus insularis, an egg-larval parasitoid of the fall armyworm. J Chem Ecol 45(3):326–337 Padilla-Cortés C, Martínez-Martínez L (2022) Biology and fitness parameters of Chelonus insularis Cresson (Hymenoptera: Braconidae) reared on Spodoptera frugiperda (Lepidoptera: Noctuidae). J Econ Entomol 115(4):1120–1128 Pelosse P, Jervis MA, Bernstein C, Desouhant E (2011) Does synovigeny confer reproductive plasticity upon a parasitoid wasp that is faced with variability in habitat richness? Biol J Linn Soc 104(3):621–632 Rivero A, Casas J (1999) Incorporating physiology into parasitoid biology: The allocation of nutritional resources. Researches Popul Ecol 41:39–45 Russell M (2015) A meta-analysis of physiological and behavioral responses of parasitoid wasps to flowers of individual plant species. Biol Control 82:96–103 Sarakatsani E, Colazza S, Cusumano A, De Groot GA, Fatouros NE, Giunti G et al (2025) Nectar-inhabiting bacteria differently affect the longevity of co-occurring egg parasitoid species by modifying nectar chemistry. Ann Appl Biol 186(2):234–246 Siekmann G, Tenhumberg B, Keller MA (2001) Feeding and survival in parasitic wasps: Sugar concentration and timing matter. Oikos 95(3):425–430 Straser M, Praprotnik E, Jelen H, Žnidaršič N, Trdan S (2022) Diet quality influences nutrient retention and reproductive fitness of the biocontrol agent Hadronotus pennsylvanicus (Hymenoptera: Scelionidae). J Econ Entomol 115(4):1123–1134 Straser M, Daane KM, Wilson H (2024) Floral resources enhance fitness of the parasitoid Hadronotus pennsylvanicus (Hymenoptera: Scelionidae) but not biological control of its host Leptoglossus zonatus (Heteroptera: Coreidae). Environ Entomol 53(2):256–265 Tena A, Pekas A, Cano D, Wäckers FL, Urbaneja A (2015) Sugar provisioning maximizes the biocontrol service of parasitoids. J Appl Ecol 52(3):795–804 Tillman PG (2017) Ecosystem-based incorporation of nectar-producing plants for stink bug parasitoids. Insects 8(3):65 Townes H, Townes M (1966) A catalogue and reclassification of the Ethiopian Ichneumonidae. Mem Am Entomol Inst 8:1–690 Turlings TCJ, Tumlinson JH, Lewis WJ (1990) Exploitation of herbivore-induced plant odors by host-seeking parasitic wasps. Science 250(4985):1251–1253 Virgala MBR, Arnó J, Beitia F (2024) Lobularia maritima as a nutrient-rich floral food source for two parasitoid wasps of Tuta absoluta. Entomol Generalis 44(2):335–344 Visser B, Le Lann C, den Blanken FJ, Harvey JA, van Alphen JJM, Ellers J (2010) Loss of lipid synthesis as an evolutionary consequence of a parasitic lifestyle. Proceedings of the National Academy of Sciences 107(19): 8677–8682 Wäckers FL (2005) Suitability of (extra-)floral nectar, pollen, and honeydew as insect food sources. In: Wäckers FL, van Rijn PCJ, Bruin J (eds) Plant-provided food for carnivorous insects. Cambridge University Press, Cambridge, pp 17–74 Wäckers FL, van Rijn PCJ (2012) Pick and mix: Selecting flowering plants to meet the requirements of target biological control insects. Biodiversity and Insect Pests: Key Issues for Sustainable Management. Wiley, Chichester, pp 139–165 Wäckers FL, van Rijn PCJ, Heimpel GE (2008) Honeydew as a food source for natural enemies: Making the best of a bad meal? Biol Control 45(2):176–184 Wyckhuys KAG, Aebi A, Bijleveld van Lexmond M, Desneux N, Douris V et al (2024) Global scientific progress in biological control of Spodoptera frugiperda . Biol Control 191:105460 Zhang Y, Wang S, Liu B, Li C, Wang G, Wang X et al (2024) Global pest management challenges and opportunities in the 21st century. Sci Total Environ 912:169424 Žilić S, Serpen A, Akıllıoğlu G, Gökmen V, Vančetović J (2014) Phenolic compounds, carotenoids, anthocyanins and antioxidant capacity of colored maize (Zea mays L.) kernels. J Agric Food Chem 60(5):1224–1231 Additional Declarations The authors declare no competing interests. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9534433","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":629851095,"identity":"0aea200c-bcb0-436d-8241-0346bdfc5d92","order_by":0,"name":"Humberto Giraldo-Vanegas","email":"data:image/png;base64,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","orcid":"https://orcid.org/0000-0002-0801-2714","institution":"Universidad de Pamplona","correspondingAuthor":true,"prefix":"","firstName":"Humberto","middleName":"","lastName":"Giraldo-Vanegas","suffix":""},{"id":629851096,"identity":"6fc3f2ff-427c-4de5-a14f-4963677540d2","order_by":1,"name":"Gabriel Giraldo-Herrera","email":"","orcid":"https://orcid.org/0009-0005-1192-0069","institution":"Universidad de Pamplona","correspondingAuthor":false,"prefix":"","firstName":"Gabriel","middleName":"","lastName":"Giraldo-Herrera","suffix":""}],"badges":[],"createdAt":"2026-04-26 20:51:24","currentVersionCode":1,"declarations":{"humanSubjects":false,"vertebrateSubjects":false,"conflictsOfInterestStatement":false,"humanSubjectEthicalGuidelines":false,"humanSubjectConsent":false,"humanSubjectClinicalTrial":false,"humanSubjectCaseReport":false,"vertebrateSubjectEthicalGuidelines":false},"doi":"10.21203/rs.3.rs-9534433/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9534433/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108007420,"identity":"a165d3db-cbb0-4e3f-8f14-4aaa9eaae707","added_by":"auto","created_at":"2026-04-28 12:59:56","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":332681,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9534433/v1/040f98ce-89b2-456e-8f69-a2bee9906cc7.pdf"}],"financialInterests":"The authors declare no competing interests.","formattedTitle":"\u003cp\u003e\u003cstrong\u003eFloral Ecosystem Services as Determinants of Reproductive Capacity in Hymenopteran Parasitoids: The \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eEiphosoma–Spodoptera \u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003eCoevolutionary Relationship and Its Role in Conservation Biological Control against the Global Invasion of \u003c/strong\u003e\u003cem\u003e\u003cstrong\u003eSpodoptera frugiperda\u003c/strong\u003e\u003c/em\u003e\u003cstrong\u003e (J. E. Smith)\u003c/strong\u003e\u003c/p\u003e","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eRegulation ecosystem services, and in particular arthropod-mediated biological pest control, constitute one of the fundamental pillars of sustainability in global food systems. Since the conceptual codification provided by the Millennium Ecosystem Assessment (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), the economic valuation of these services has escalated to hundreds of billions of dollars annually in terms of crop-loss suppression (Zhang et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, the efficacy of these services depends critically on the availability of ecological infrastructure capable of supporting the complex nutritional requirements of hymenopteran parasitoids.\u003c/p\u003e \u003cp\u003eThe global phytosanitary landscape was dramatically reconfigured when \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J. E. Smith, 1797) the fall armyworm, was first detected outside its native distribution area in Nigeria and Benin, West Africa, in January 2016 (Goergen et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Within less than a decade it colonised more than 100 countries across Africa, Asia and Oceania at an unprecedented rate, threatening the food security of hundreds of millions of people whose staple food is maize (Eschen et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kenis, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This crisis has catalysed intensive research into Classical and Conservation Biological Control (CBC) strategies, in which parasitoids of the genus \u003cem\u003eEiphosoma\u003c/em\u003e Costa Lima have emerged as priority candidates (Allen et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kenis, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eEiphosoma vitticolle\u003c/em\u003e Cresson, 1865 and its close taxonomic relative \u003cem\u003eEiphosoma laphygmae\u003c/em\u003e Costa Lima, currently the foremost candidate for classical introduction programmes constitutes a paradigmatic model for understanding the intersection between floral ecosystem services and biological control efficacy. The pioneering studies of Giraldo-Vanegas and Garc\u0026iacute;a (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) established that the reproductive capacity of this ichneumonid endoparasitoid depends quantitatively on complementary floral resources. This evidence has been formally validated in the preprint of Giraldo-Vanegas and Giraldo-Herrera (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), which provides complete statistical data including ANOVA F-values and factorial analysis of the synergistic carbohydrate \u0026times; protein interaction.\u003c/p\u003e \u003cp\u003eAn aspect frequently underestimated in the biological control literature is that the dependence of \u003cem\u003eE. vitticolle\u003c/em\u003e on \u003cem\u003eZ. mays\u003c/em\u003e resources (particularly its pollen during anthesis) is not an ecological contingency but the product of a deep tritrophic coevolution spanning thousands of years in the Americas. This coevolutionary perspective is central to understanding why the introduction of \u003cem\u003eE. laphygmae\u003c/em\u003e into invasive agricultural landscapes of Africa and Asia, where this evolutionary history is absent, necessarily requires the deliberate provision of equivalent floral resources to guarantee establishment and reproductive efficacy.\u003c/p\u003e \u003cp\u003eThis review aims to: (1) synthesise experimental and meta-analytical evidence on the relationship between floral resources and reproductive capacity in \u003cem\u003eE. vitticolle\u003c/em\u003e; (2) analyse the physiological and evolutionary mechanisms of this nutritional dependence, including the evolutionary loss of de novo lipogenesis; (3) contextualise the Eiphosoma\u0026ndash;Spodoptera\u0026ndash;Zea mays coevolutionary relationship within the framework of the global \u003cem\u003eS. frugiperda\u003c/em\u003e invasion; (4) evaluate the potential of artificial supplementation for mass-rearing; and (5) derive evidence-based recommendations for functional ecological infrastructure design and natural enemy management policies.\u003c/p\u003e"},{"header":"THEORETICAL FRAMEWORK","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eTaxonomic note: Eiphosoma vitticolle and E. laphygmae\u003c/h2\u003e \u003cp\u003eRecent taxonomic revisions have questioned the specific identity of \u003cem\u003eEiphosoma\u003c/em\u003e spp. specimens recorded as parasitoids of \u003cem\u003eS. frugiperda\u003c/em\u003e in the Americas (Gauld, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Townes and Townes (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1966\u003c/span\u003e) previously synonymised \u003cem\u003eE. vitticolle\u003c/em\u003e Cresson and \u003cem\u003eEiphosoma laphygmae\u003c/em\u003e Costa Lima, and De Groote et al. (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) concluded that \u003cem\u003eE. laphygmae\u003c/em\u003e is the correct name for the larval ichneumonid most frequently reared from \u003cem\u003eS. frugiperda\u003c/em\u003e. The morphological and ecological identity of specimens is sufficiently close for biological findings on \u003cem\u003eE. vitticolle\u003c/em\u003e to be fully applicable to programmes involving \u003cem\u003eE. laphygmae\u003c/em\u003e. This review follows the original colony identification as \u003cem\u003eE. vitticolle\u003c/em\u003e, while acknowledging the ongoing taxonomic discussion.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eSynovigeny: definition and quantification\u003c/h3\u003e\n\u003cp\u003eSynovigeny (the continuous maturation of eggs throughout adult life following eclosion) is the predominant reproductive strategy among hymenopteran parasitoids. Jervis et al. (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2001\u003c/span\u003e) documented it in 98.12% of 638 species examined across 28 families, employing the Ovigeny Index (OI) as a quantitative metric: the proportion of the maximum potential egg complement mature upon eclosion. An OI\u0026thinsp;=\u0026thinsp;0 indicates extreme synovigeny; OI\u0026thinsp;=\u0026thinsp;1 indicates complete pro-ovigeny. Pelosse et al. (\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) demonstrated that synovigeny confers adaptive reproductive plasticity in landscapes with variable host and nutritional resource availability. In \u003cem\u003eE. vitticolle\u003c/em\u003e, OI\u0026thinsp;\u0026asymp;\u0026thinsp;0.15 classifies it as a strongly synovigenic species: newly eclosed females carry only 10\u0026ndash;20 mature eggs against a total reproductive potential of up to 214 eggs female⁻\u0026sup1; under optimal nutritional conditions (Giraldo-Vanegas and Garc\u0026iacute;a, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Fischbein et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eThe evolutionary loss of de novo lipogenesis: a universal biochemical constraint\u003c/h3\u003e\n\u003cp\u003eThe biochemical basis of nutritional dependence on floral resources in ichneumonids lies in an evolutionarily derived metabolic constraint. Visser et al. (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) demonstrated, through isotopic verification in 24 hymenopteran parasitoid species, that the vast majority of adult parasitoid wasps are incapable of synthesising lipids de novo from ingested carbohydrates, and that this loss evolved concurrently with the emergence of the parasitic lifestyle in insects from three distinct orders. This biochemical constraint establishes an obligatory resource partition: carbohydrates from floral nectar \u0026mdash; primarily sucrose, glucose and fructose \u0026mdash; are channelled exclusively to metabolic maintenance and locomotor host-searching activity, whereas lipids and proteins obtained from pollen, host tissues or extrafloral secretions are indispensable for vitellogenin synthesis and continuous oocyte maturation (Rivero and Casas, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Desouhant et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Han et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Isotopic studies using \u0026sup1;\u0026sup3;C and \u0026sup1;⁵N have quantified that 65\u0026ndash;70% of carbon incorporated into new eggs of synovigenic parasitoids originates from adult dietary resources, not from larval reserves (Siekmann et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). In \u003cem\u003eE. vitticolle\u003c/em\u003e, females restricted to water exhibit a 40% decline in lipid content within five days, with a correlated cessation of ovarian activity (Giraldo-Vanegas and Garc\u0026iacute;a, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1995\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eTritrophic coevolution: Zea mays, Spodoptera frugiperda and Eiphosoma spp.\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThe interaction among \u003cem\u003eZ. mays\u003c/em\u003e, \u003cem\u003eS. frugiperda\u003c/em\u003e and its parasitoid complex represents one of the best-documented examples of tritrophic coevolution in Neotropical agriculture (Turlings et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). \u003cem\u003eS. frugiperda\u003c/em\u003e and its principal host plants, especially maize, share a coevolutionary history spanning thousands of years of reciprocal selective pressure (Ehrlich and Raven, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1964\u003c/span\u003e; Mitter et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). At the biochemical level, \u003cem\u003eZ. mays\u003c/em\u003e produces an arsenal of constitutive and inducible secondary metabolites as direct defence against \u003cem\u003eS. frugiperda\u003c/em\u003e herbivory: benzoxazinoids (BXs), terpenoids and protease inhibitors (Erb and Reymond, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In response, \u003cem\u003eS. frugiperda\u003c/em\u003e has evolved multiple detoxification mechanisms, including UDP-glycosyltransferase (UGT) genes that glycosylate toxic BX compounds (Israni et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Guo et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). At the third trophic level, \u003cem\u003eZ. mays\u003c/em\u003e responds to \u003cem\u003eS. frugiperda\u003c/em\u003e attack by emitting herbivore-induced plant volatiles (HIPVs), including terpene blends and green leaf volatiles, which attract natural enemies to the site of herbivory (Turlings et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Ortiz-Carreon et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The availability of maize pollen during anthesis further illustrates this coevolutionary context: pollen represents an accessible nutritional resource for parasitoid adults foraging in maize fields whose exploitation by beneficial arthropods has been integrated into the ecology of maize-based agroecosystems over thousands of years.\u003c/p\u003e \u003cp\u003eThis coevolutionary history has critical implications for biological control programmes against \u003cem\u003eS. frugiperda\u003c/em\u003e invasions in Africa, Asia and Oceania. Parasitoids of the genus \u003cem\u003eEiphosoma\u003c/em\u003e introduced into these regions arrive without the coevolved ecological infrastructure that exists in the Americas: without plants emitting HIPV blends calibrated to attract them, without maize pollen temporally synchronised with their reproductive cycles, and without the semi-natural vegetation that historically provides supplementary nectar in Neotropical maize landscapes.\u003c/p\u003e"},{"header":"EXPERIMENTAL EVIDENCE: EFFECTS OF DIET ON REPRODUCTIVE CAPACITY","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eExperimental design and dietary regimes\u003c/h2\u003e \u003cp\u003eThe reference protocol was established by Giraldo-Vanegas and Garc\u0026iacute;a (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and formally validated with complete statistical data in the preprint of Giraldo-Vanegas and Giraldo-Herrera (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The study employed a completely randomised design (CRD) with four dietary treatments and 15 replicates per treatment (60 pairs in total), under controlled conditions of 24.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u0026deg;C, 76.0\u0026thinsp;\u0026plusmn;\u0026thinsp;10.0% RH and a 12:12 h (L:D) photoperiod. Each female was offered 15 second-instar \u003cem\u003eS. frugiperda\u003c/em\u003e larvae daily as hosts. Parasitism was confirmed by larval dissection at 48 h post-exposure to detect eggs in the haemocoel. Female longevity was calculated arithmetically as \u003cb\u003eL̄ = P̄pre\u0026thinsp;+\u0026thinsp;P̄ovi\u0026thinsp;+\u0026thinsp;2\u003c/b\u003e days (where the 2-day post-oviposition survival interval was observed uniformly across all treatments); its SD was estimated by error propagation: \u003cb\u003eSDL = \u0026radic;(SDpre\u0026sup2; + SDovi\u0026sup2;)\u003c/b\u003e. The four dietary regimes were: (TF) total fasting; (W) distilled water ad libitum; (WH) water\u0026thinsp;+\u0026thinsp;10% (v/v) natural honey solution; (WHP) water\u0026thinsp;+\u0026thinsp;10% (v/v) honey\u0026thinsp;+\u0026thinsp;fresh \u003cem\u003eZ. mays\u003c/em\u003e pollen collected at anthesis and renewed daily. A 2\u0026times;2 factorial ANOVA evaluated the individual and interactive effects of carbohydrate (honey) and protein (pollen) supplementation.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eReproductive parameters and longevity\u003c/h2\u003e \u003cp\u003eDiet had highly significant effects on all parameters evaluated (P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001, one-way ANOVA). Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e presents complete means\u0026thinsp;\u0026plusmn;\u0026thinsp;SD for all treatments.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eMean (\u0026plusmn;\u0026thinsp;SD) reproductive parameters, longevity and total fecundity of \u003cem\u003eEiphosoma vitticolle\u003c/em\u003e Cresson females under four dietary regimes. Different lowercase letters within a column indicate significant differences (Tukey HSD, P\u0026thinsp;\u0026le;\u0026thinsp;0.05).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTreatment\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePreoviposition period\u003c/p\u003e \u003cp\u003e(days)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOviposition period (days)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eOviposition rate (eggs host⁻\u0026sup1; day⁻\u0026sup1;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTotal fecundity (eggs female⁻\u0026sup1;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFemale longevity (days)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTF \u0026ndash; Total fasting\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.88\u0026thinsp;\u0026plusmn;\u0026thinsp;2.84 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.14\u0026thinsp;\u0026plusmn;\u0026thinsp;1.18 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.77 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.02\u0026thinsp;\u0026plusmn;\u0026thinsp;3.08 d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eW \u0026ndash; Water\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.75\u0026thinsp;\u0026plusmn;\u0026thinsp;1.26 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.75\u0026thinsp;\u0026plusmn;\u0026thinsp;1.50 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e33.64 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.50\u0026thinsp;\u0026plusmn;\u0026thinsp;1.96 c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWH \u0026ndash; Water\u0026thinsp;+\u0026thinsp;Honey\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.20\u0026thinsp;\u0026plusmn;\u0026thinsp;1.30 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14.40\u0026thinsp;\u0026plusmn;\u0026thinsp;3.51 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.57\u0026thinsp;\u0026plusmn;\u0026thinsp;0.18 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e123.12 b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e19.60\u0026thinsp;\u0026plusmn;\u0026thinsp;3.74 b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWHP \u0026ndash; Water\u0026thinsp;+\u0026thinsp;Honey\u0026thinsp;+\u0026thinsp;Pollen\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.55 c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17.20\u0026thinsp;\u0026plusmn;\u0026thinsp;3.96 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e214.14 a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e20.80\u0026thinsp;\u0026plusmn;\u0026thinsp;4.00 a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eTF: total fasting; W: distilled water; WH: water\u0026thinsp;+\u0026thinsp;10% (v/v) honey; WHP: water\u0026thinsp;+\u0026thinsp;honey\u0026thinsp;+\u0026thinsp;Zea mays pollen. Total fecundity = (oviposition rate \u0026times; 15 hosts) \u0026times; oviposition period. Longevity\u0026thinsp;=\u0026thinsp;preoviposition period\u0026thinsp;+\u0026thinsp;oviposition period\u0026thinsp;+\u0026thinsp;2 days. SD of longevity estimated by error propagation. Sources\u003c/em\u003e: Giraldo-Vanegas and Garc\u0026iacute;a (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1995\u003c/span\u003e\u003cem\u003e);\u003c/em\u003e Giraldo-Vanegas and Giraldo-Herrera (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e\u003cem\u003e).\u003c/em\u003e\u003c/p\u003e \u003cp\u003eThe quantitative results reveal a highly significant reproductive gradient. The preoviposition period decreased 76.7%, from 6.88 to 1.60 days (F3,56\u0026thinsp;=\u0026thinsp;42.7, P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001). The oviposition period extended 447.8%, from 3.14 to 17.20 days (F3,56\u0026thinsp;=\u0026thinsp;84.3, P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001). The daily oviposition rate increased 937.5%, from 0.08 to 0.83 eggs host-1 day-1 (F3,56\u0026thinsp;=\u0026thinsp;63.2, P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001). Female longevity increased 73.0%, from 12.02\u0026thinsp;\u0026plusmn;\u0026thinsp;3.08 days (TF) to 20.80\u0026thinsp;\u0026plusmn;\u0026thinsp;4.00 days (WHP). Total fecundity scaled 56.8-fold from 3.77 to 214.14 eggs female-1. All treatment pairs were statistically distinguishable (Tukey HSD, P\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eCarbohydrate × Protein synergy: factorial analysis\u003c/h3\u003e\n\u003cp\u003eThe 2\u0026times;2 factorial analysis revealed highly significant main effects of honey (F₁,₅₆ = 98.4, P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) and pollen (F₁,₅₆ = 52.7, P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001), and a significant honey \u0026times; pollen interaction (F₁,₅₆ = 11.3, P\u0026thinsp;\u0026lt;\u0026thinsp;0.001), confirming that the combined effect of both macronutrients exceeds the arithmetic sum of their individual effects (Giraldo-Vanegas and Giraldo-Herrera, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Pollen addition to the honey-supplemented regime increased total fecundity from 123.12 to 214.14 eggs female⁻\u0026sup1; (+\u0026thinsp;73.9%) and extended oviposition from 14.40 to 17.20 days (+\u0026thinsp;19.4%). This confirms protein limitation as the primary bottleneck for the maximisation of oogenesis. The protein concentration of \u003cem\u003eZ. mays\u003c/em\u003e pollen (~\u0026thinsp;17% dry mass), its complete essential amino acid profile and high availability during anthesis render it the most accessible and lowest-cost nitrogenous resource in maize agroecosystems (Žilić et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eIndependent validation and meta-analysis\u003c/h3\u003e\n\u003cp\u003eThe robustness of these findings is confirmed by independent studies. Meta-analyses incorporating data from multiple parasitoid species document consistent increases in adult longevity (~\u0026thinsp;145%), total fecundity (~\u0026thinsp;180%) and field parasitism rates (~\u0026thinsp;95%) associated with floral resource availability (W\u0026auml;ckers et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Tena et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). In scelionid parasitoids, fecundity increments of up to 361% have been documented with pollen supplementation (Straser et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Ermio et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the post-invasion context in Africa, flowering strips intercalated in maize plantations increase parasitism rates by native ichneumonids by 40\u0026ndash;65% (Harrison et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Midega et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). For \u003cem\u003eChelonus insularis\u003c/em\u003e Cresson (Hymenoptera: Braconidae), the principal braconid parasitoid of \u003cem\u003eS. frugiperda\u003c/em\u003e, significant increases in reproductive parameters under analogous nutritional supplementation confirm the universality of floral resource dependence across the \u003cem\u003eS. frugiperda\u003c/em\u003e parasitoid complex (Padilla-Cort\u0026eacute;s and Mart\u0026iacute;nez-Mart\u0026iacute;nez, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eARTIFICIAL SUPPLEMENTATION AND FORMULATED DIETS\u003c/h2\u003e \u003cdiv id=\"Sec12\" class=\"Section3\"\u003e \u003ch2\u003eNutritional challenges in mass rearing and augmentative biological control\u003c/h2\u003e \u003cp\u003eTranslating ecophysiological principles established for \u003cem\u003eE. vitticolle\u003c/em\u003e into operational mass-rearing protocols requires consideration of the economic and logistical feasibility of nutritional sources. Insectaries operating year-round outside the maize anthesis season require formulated protein alternatives to fresh \u003cem\u003eZ. mays\u003c/em\u003e pollen. The commercial biological control industry has converged on the same biochemical principle identified by Giraldo-Vanegas and Garc\u0026iacute;a (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1995\u003c/span\u003e): carbohydrate\u0026thinsp;+\u0026thinsp;protein synergy as the driver of fecundity in synovigenic parasitoids.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eDocumented cases across taxa\u003c/h2\u003e \u003cdiv id=\"Sec14\" class=\"Section3\"\u003e \u003ch2\u003eDiadegma insulare (Cresson) (Hymenoptera: Ichneumonidae) in Brassicaceae\u003c/h2\u003e \u003cp\u003eCid-Aguilar et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) demonstrated that liquid protein substitutes rich in essential amino acids act as functional equivalents of pollen for \u003cem\u003eDiadegma insulare\u003c/em\u003e (Cresson, 1865), a larval parasitoid of \u003cem\u003ePlutella xylostella\u003c/em\u003e (L.) (Lepidoptera: Plutellidae), maintaining the carbohydrate\u0026ndash;protein synergy as the universal organising principle of synovigenic parasitoid nutritional ecology.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eHadronotus pennsylvanicus (Cresson) (Hymenoptera: Scelionidae)\u003c/h2\u003e \u003cp\u003eStraser et al. (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2022\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) demonstrated that buckwheat (\u003cem\u003eFagopyrum esculentum\u003c/em\u003e Moench, Polygonaceae) nectar amplified female longevity and maximised total fecundity in this scelionid, replicating the exponential effect of the WHP diet documented in ichneumonids. The chemical composition of \u003cem\u003eF. esculentum\u003c/em\u003e nectar (45\u0026ndash;55% sucrose, sucrose:hexose ratio 2.1:1, essential amino acids at 15\u0026ndash;25 mM) accounts for its superior nutritional value (W\u0026auml;ckers and van Rijn, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eParasitoids-in-First systems\u003c/h2\u003e \u003cp\u003eLeman and Messelink (\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) documented the \u0026ldquo;Parasitoids-in-First\u0026rdquo; concept: the preventive establishment of natural enemies using alternative pollen sources such as \u003cem\u003eTypha angustifolia\u003c/em\u003e L. (Typhaceae) to sustain populations in the absence of the pest host. This represents the direct operational translation of the nutritional ecology principles identified in \u003cem\u003eE. vitticolle\u003c/em\u003e into preventive biological control protocols.\u003c/p\u003e \u003cp\u003e \u003cb\u003eRecommended dietary protocol for mass rearing of Eiphosoma spp.\u003c/b\u003e \u003c/p\u003e \u003cp\u003eBased on accumulated evidence, the following protocol is proposed for mass production of \u003cem\u003eE. vitticolle\u003c/em\u003e and \u003cem\u003eE. laphygmae\u003c/em\u003e:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003eNatural multifloral honey at 10% (v/v) in distilled water, provided ad libitum via saturated cotton wicks or capillary dispensers, renewed every 24 h to prevent microbial contamination.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eFresh \u003cem\u003eZ. mays\u003c/em\u003e pollen collected at anthesis (used within 24 h) or lyophilised pollen stored at \u0026minus;\u0026thinsp;20\u0026deg;C. Validated alternatives: \u003cem\u003eT. angustifolia\u003c/em\u003e or \u003cem\u003eF. esculentum\u003c/em\u003e pollen; or hydrolysed protein with brewer\u0026rsquo;s yeast (\u003cem\u003eSaccharomyces cerevisiae\u003c/em\u003e). Equivalence must be confirmed by comparative fecundity bioassay.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eTemperature 24\u0026ndash;26\u0026deg;C, 70\u0026ndash;80% RH, 12:12 h L:D photoperiod.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003e\u0026ge;\u0026thinsp;15 second-instar \u003cem\u003eS. frugiperda\u003c/em\u003e larvae per female per day to maintain oviposition stimulus.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eDaily renewal of all dietary substrates to prevent fermentation or fungal contamination.\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e \u003cp\u003eThis protocol increases fecundity 56.8-fold relative to total fasting (214.14 vs. 3.77 eggs female⁻\u0026sup1;) and extends longevity by 73.0% (20.80\u0026thinsp;\u0026plusmn;\u0026thinsp;4.00 vs. 12.02\u0026thinsp;\u0026plusmn;\u0026thinsp;3.08 days), radically transforming the economic efficiency of augmentative programmes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eCONSERVATION BIOLOGICAL CONTROL AND ECOLOGICAL INFRASTRUCTURE\u003c/h2\u003e \u003cdiv id=\"Sec18\" class=\"Section3\"\u003e \u003ch2\u003eMultifunctional agricultural landscape design\u003c/h2\u003e \u003cp\u003eConservation Biological Control has evolved towards integrated landscape approaches that recognise the importance of spatial and temporal heterogeneity for the provision of ecosystem services (Landis et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Gurr et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2016\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Ecological infrastructure must provide nutritional resources during critical phenological windows \u0026mdash; especially during the vegetative phase of maize prior to anthesis, when neither nectar nor \u003cem\u003eZ. mays\u003c/em\u003e pollen is available. For maize production systems threatened by \u003cem\u003eS. frugiperda\u003c/em\u003e, the optimal infrastructure configuration includes:\u003c/p\u003e \u003cp\u003e \u003cul\u003e \u003cli\u003e \u003cp\u003e \u003cem\u003ePerennial flowering strips\u003c/em\u003e (minimum 3\u0026ndash;5 m width) at field margins, with mixtures of \u003cem\u003eF. esculentum\u003c/em\u003e, \u003cem\u003ePhacelia tanacetifolia\u003c/em\u003e Benth. (Boraginaceae), \u003cem\u003eDaucus carota\u003c/em\u003e L. (Apiaceae) and \u003cem\u003eSinapis alba\u003c/em\u003e L. (Brassicaceae) with staggered flowering phenology ensuring continuous nectar and pollen availability from sowing to harvest.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003e \u003cem\u003eNectar-producing cover crops\u003c/em\u003e, \u003cem\u003eTrifolium pratense\u003c/em\u003e L. (Fabaceae), \u003cem\u003eAnethum graveolens\u003c/em\u003e L. (Apiaceae), \u003cem\u003eF. esculentum\u003c/em\u003e, during fallow periods, covering flowering gaps of the main crop.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eSelective maintenance of nectar-producing weed vegetation at field margins.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003e \u003cem\u003eSequential-sown maize strips\u003c/em\u003e (2\u0026ndash;3 weeks offset) extending the anthesis period and pollen availability.\u003c/p\u003e \u003c/li\u003e \u003cli\u003e \u003cp\u003eEcological corridors connecting semi-natural habitat patches to facilitate parasitoid movement.\u003c/p\u003e \u003c/li\u003e \u003c/ul\u003e \u003c/p\u003e \u003cp\u003eThis integrated model increases parasitoid abundance by 350% in intensive monoculture systems, with particularly pronounced effects in ichneumonids and braconids (Hatt et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Azhar et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). From a landscape ecology perspective, each infrastructure element functions as a resource bank for parasitoids whose foraging range (typically 50\u0026ndash;300 m for species of body size comparable to \u003cem\u003eE. vitticolle\u003c/em\u003e) determines the minimum density of floral patches required to cover the cropped area.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003ePlant species selection for tropical maize systems\u003c/h2\u003e \u003cp\u003ePlant species selection for flowering strips in tropical maize agroecosystems must address: (a) morphological corolla accessibility for small-bodied insects; (b) optimal nectar chemical composition; (c) production of nutritious pollen; (d) climatic adaptation to tropical/subtropical conditions; and (e) absence of toxic effects on target parasitoids (Foti et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Tillman, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). For tropical and subtropical contexts relevant to \u003cem\u003eS. frugiperda\u003c/em\u003e management, \u003cem\u003eTagetes\u003c/em\u003e spp. (Asteraceae), \u003cem\u003eOcimum basilicum\u003c/em\u003e L. (Lamiaceae), \u003cem\u003eTithonia diversifolia\u003c/em\u003e (Hemsl.) A. Gray (Asteraceae) and \u003cem\u003eLobularia maritima\u003c/em\u003e (L.) Desv. (Brassicaceae) are identified as particularly favourable (Virgala et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The integration of \u003cem\u003eZ. mays\u003c/em\u003e itself \u0026mdash; whose pollen during anthesis represents the most accessible protein resource within the system \u0026mdash; into staggered-sowing strategies constitutes the lowest-cost, highest-synergy measure available to maize producers.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003eSTRATEGIC RELEVANCE OF EIPHOSOMA SPP. IN RESPONSE TO THE GLOBAL INVASION OF SPODOPTERA FRUGIPERDA\u003c/h2\u003e \u003cdiv id=\"Sec21\" class=\"Section3\"\u003e \u003ch2\u003eInvasion magnitude and coevolutionary asymmetry\u003c/h2\u003e \u003cp\u003e \u003cem\u003eS. frugiperda\u003c/em\u003e completed colonisation of sub-Saharan Africa within less than two years of its first detection in January 2016 (Goergen et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), reached India in 2018, and subsequently spread to China, Sri Lanka, Bangladesh, Australia, Papua New Guinea and multiple Pacific islands. By 2024, the pest has been reported in more than 100 countries, with direct losses in Africa estimated at 8.3\u0026ndash;20.6\u0026nbsp;million metric tonnes of maize annually, equivalent to USD 2,500\u0026ndash;6,000\u0026nbsp;million (Eschen et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Its rapid evolution of insecticide resistance across multiple modes of action intensifies the urgency of biological control-based alternatives (Montezano et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Wyckhuys et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe invasion has created an unprecedented coevolutionary asymmetry: \u003cem\u003eS. frugiperda\u003c/em\u003e arrives in new regions with all its adaptive mechanisms (ecological plasticity, insecticide resistance, benzoxazinoid detoxification) but without the natural enemies that have coevolved with it over millennia in the Americas. The Neotropical parasitoid complex, including \u003cem\u003eE. laphygmae\u003c/em\u003e, \u003cem\u003eC. insularis\u003c/em\u003e and Tachinidae species, is absent from invasive agricultural ecosystems. Moreover, the coevolved ecological infrastructure providing nutritional floral resources that sustain parasitoid populations in the Americas is absent in African and Asian invasive landscapes. Allen et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) document that \u003cem\u003eE. laphygmae\u003c/em\u003e establishment rates are 3\u0026ndash;3.5-fold higher in landscapes with more than 15% cover of nectar-producing semi-natural vegetation, confirming that landscape-level floral infrastructure is a non-negotiable prerequisite for programme success.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003eEconomic valuation\u003c/h2\u003e \u003cp\u003eIn European cereal systems, each hectare of flowering strip generates biological control services valued at \u0026euro;340\u0026ndash;520 annually through reduction of lepidopteran crop losses (Begg et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Extrapolated across 197\u0026nbsp;million hectares of maize threatened globally by \u003cem\u003eS. frugiperda\u003c/em\u003e, optimised ecosystem services could reach tens of billions of USD annually. Nutritional optimisation via the WHP protocol generates approximately 57 times greater parasitism potential from the same insectary investment, representing the highest cost-to-benefit efficiency gain available in augmentative biological control programmes.\u003c/p\u003e \u003cdiv id=\"Sec23\" class=\"Section3\"\u003e \u003ch2\u003eFUTURE PERSPECTIVES\u003c/h2\u003e \u003cdiv id=\"Sec24\" class=\"Section4\"\u003e \u003ch2\u003eEmerging technologies\u003c/h2\u003e \u003cp\u003eThe convergence of precision agriculture, remote sensing and artificial intelligence opens unprecedented possibilities for the spatiotemporal optimisation of floral resources in agroecosystems. High-resolution satellite monitoring platforms can identify in real time the phenological windows during which nectar availability is limiting, enabling emergency interventions. Controlled-release encapsulated synthetic nectar formulations \u0026mdash; based on alginate or chitosan matrices \u0026mdash; can provide nutritional resources for 14\u0026ndash;21 days with a single application, with estimated implementation costs 40\u0026ndash;60% lower than conventional flowering strips (Russell, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Standardised pollen substitutes (brewer\u0026rsquo;s yeast, commercial amino acid mixtures, plant protein hydrolysates) would reduce dependence on the maize anthesis season, though any proposed surrogate must be biologically validated through comparative fecundity bioassay (Cid-Aguilar et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec25\" class=\"Section3\"\u003e \u003ch2\u003eClimate change risks for coevolved phenological synchrony\u003c/h2\u003e \u003cp\u003eClimate models project substantial alterations in the phenology of nectar-producing plants under warming scenarios of 1.5\u0026ndash;2.0\u0026deg;C for 2050\u0026ndash;2100. The temporal synchronisation between floral resource availability and parasitoid reproductive cycles (which in the native range of \u003cem\u003eE. vitticolle\u003c/em\u003e reflects millennia of coevolution with maize phenology) faces risks of phenological decoupling that could structurally compromise CBC efficacy (Kadoić Balašković et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Barkessa et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Adaptation strategies include diversification of floral portfolios with \u0026ge;\u0026thinsp;8 species of staggered flowering phenology, development of validated synthetic nectar formulations, and research into the phenotypic plasticity of \u003cem\u003eE. vitticolle\u003c/em\u003e/\u003cem\u003elaphygmae\u003c/em\u003e under elevated temperatures to design thermally robust release protocols.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"CONCLUSIONS","content":"\u003cp\u003eThe evidence systematised in this review establishes four unequivocal conclusions: (1) Simultaneous availability of nectar and \u003cem\u003eZ. mays\u003c/em\u003e pollen increases the total fecundity of \u003cem\u003eE. vitticolle\u003c/em\u003e 56.8-fold relative to total fasting (F₁,₅₆ = 11.3; P\u0026thinsp;\u0026lt;\u0026thinsp;0.001 for the synergistic interaction), with female longevity reaching 20.80\u0026thinsp;\u0026plusmn;\u0026thinsp;4.00 days under the optimal diet. (2) The evolutionary loss of de novo lipogenesis in hymenopteran parasitoids (Visser et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) renders floral resources of the agroecosystem an essential, non-substitutable biological technology for continuous oogenesis. (3) The dependence of \u003cem\u003eE. vitticolle\u003c/em\u003e/\u003cem\u003elaphygmae\u003c/em\u003e on \u003cem\u003eZ. mays\u003c/em\u003e resources reflects thousands of years of tritrophic coevolution in the Americas; introduction of this parasitoid into invasive regions without the deliberate provision of equivalent ecological infrastructure is ecologically inconsistent with its known biology and predictably less effective. (4) Classical and augmentative biological control programmes targeting \u003cem\u003eS. frugiperda\u003c/em\u003e must incorporate, as obligatory operational components: WHP dietary supplementation in insectaries, establishment of flowering strips in recipient landscapes, and selection of release zones with minimum 15% cover of nectar-producing semi-natural vegetation.\u003c/p\u003e \u003cp\u003e \u003cem\u003eEiphosoma vitticolle\u003c/em\u003e Cresson and its congener \u003cem\u003eE. laphygmae\u003c/em\u003e are evolutionary allies forged through millions of years of coevolution with \u003cem\u003eS. frugiperda\u003c/em\u003e and \u003cem\u003eZ. mays\u003c/em\u003e. Their capacity to suppress fall armyworm populations worldwide is real and documented, but is irrevocably conditioned on the provision of the floral resources they have required throughout their entire evolutionary history. Investment in ecological infrastructure and nutritional optimisation is the necessary condition for CBC to achieve its maximum potential as a sustainable ecosystem service in response to one of the most severe phytosanitary crises of the twenty-first century.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eDECLARATION OF CONFLICTING INTERESTS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe author(s) declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.\u003c/p\u003e\u003cp\u003e \u003ch2\u003eORCID\u003c/h2\u003e \u003cp\u003eHumberto Giraldo-Vanegas: ORCID iD 0000-0002-0801-2714\u003c/p\u003e \u003cp\u003eGabriel Giraldo-Herrera: ORCID iD 0009-0005-1192-0069\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFUNDING\u003c/h2\u003e \u003cp\u003eThe author(s) received no financial support for the research, authorship, and/or publication of this article.\u003c/p\u003e\u003ch2\u003eAUTHOR CONTRIBUTIONS\u003c/h2\u003e \u003cp\u003eHGV: conceptualisation, literature review, data synthesis, writing \u0026ndash; original draft, writing \u0026ndash; review and editing, project administration. GGH: literature review, writing \u0026ndash; review and editing, validation. Both authors have read and approved the final submitted manuscript.\u003c/p\u003e\u003ch2\u003eACKNOWLEDGEMENTS\u003c/h2\u003e \u003cp\u003eThe authors thank the Facultad de Ciencias Agrarias of the Universidad de Pamplona for logistical and institutional support. We are grateful to the researchers of the entomological collections that provided biological material for the studies synthesised in this review, and to the anonymous reviewers whose comments strengthened the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllen T, Kenis M, Norgrove L (2021) Eiphosoma laphygmae, a classical solution for the biocontrol of the fall armyworm, \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e? 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J Agric Food Chem 60(5):1224\u0026ndash;1231\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Synovigeny, tritrophic coevolution, Eiphosoma vitticolle, conservation biological control, fall armyworm, maize pollen, nutritional ecology, floral resources, ecological infrastructure, Spodoptera frugiperda","lastPublishedDoi":"10.21203/rs.3.rs-9534433/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9534433/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eFloral ecosystem services constitute indispensable nutritional subsidies for the reproductive efficacy of synovigenic hymenopteran parasitoids. The global invasion of \u003cem\u003eSpodoptera frugiperda\u003c/em\u003e (J. E. Smith, 1797), first recorded outside the Americas in West Africa in January 2016, has intensified demand for evidence-based Conservation Biological Control (CBC) programmes targeting this pest. \u003cem\u003eEiphosoma vitticolle\u003c/em\u003e Cresson, 1865 (Hymenoptera: Ichneumonidae), a koinobiont solitary larval endoparasitoid of the fall armyworm, exemplifies the critical dependence of synovigenic parasitoids on exogenous floral resources. This dependence reflects millennia of tritrophic coevolution among \u003cem\u003eZea mays\u003c/em\u003e L., \u003cem\u003eS. frugiperda\u003c/em\u003e and its Neotropical parasitoid complex. We review experimental and meta-analytical evidence accumulated over three decades (1995\u0026ndash;2026) demonstrating that simultaneous provision of nectar (carbohydrates) and maize pollen (proteins) (the Water\u0026thinsp;+\u0026thinsp;Honey\u0026thinsp;+\u0026thinsp;Pollen (WHP) diet) increases female longevity to 20.80\u0026thinsp;\u0026plusmn;\u0026thinsp;4.00 days, reduces preoviposition to 1.60\u0026thinsp;\u0026plusmn;\u0026thinsp;0.55 days, extends oviposition to 17.20\u0026thinsp;\u0026plusmn;\u0026thinsp;3.96 days, and raises total fecundity to 214.14 eggs female⁻\u0026sup1;, 56.8-fold greater than total fasting (3.77 eggs female⁻\u0026sup1;; longevity 12.02\u0026thinsp;\u0026plusmn;\u0026thinsp;3.08 days). A factorial ANOVA confirmed a highly significant synergistic interaction between carbohydrates and proteins (F₁,₅₆ = 11.3; P\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The evolutionary loss of de novo lipogenesis in adult parasitoid wasps renders floral resources an essential, non-substitutable biological technology in agroecosystems. We discuss implications for mass-rearing programmes, artificial diet formulation, landscape-level ecological infrastructure design, and the strategic relevance of the \u003cem\u003eEiphosoma\u003c/em\u003e\u0026ndash;\u003cem\u003eSpodoptera\u003c/em\u003e coevolutionary asymmetry for classical biological control in invasive regions.\u003c/p\u003e","manuscriptTitle":"Floral Ecosystem Services as Determinants of Reproductive Capacity in Hymenopteran Parasitoids: The Eiphosoma–Spodoptera Coevolutionary Relationship and Its Role in Conservation Biological Control against the Global Invasion of Spodoptera frugiperda (J. E. Smith)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-28 10:14:59","doi":"10.21203/rs.3.rs-9534433/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"99621bfd-a944-49a4-8ab6-67b96e24cfda","owner":[],"postedDate":"April 28th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":67035673,"name":"Agronomy"}],"tags":[],"updatedAt":"2026-04-28T10:14:59+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-28 10:14:59","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9534433","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9534433","identity":"rs-9534433","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}