Plant architecture shapes arthropod communities and mediates indirect defense in maize landraces

Plant architecture shapes arthropod communities and mediates indirect defense in maize landraces | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Plant architecture shapes arthropod communities and mediates indirect defense in maize landraces Julio Cesar Ahuatzin-Hernández, Aldo Daniel Chan-Arjona, Alexis Lamz-Piedra, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8456967/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 Maize ( Zea mays L.) is a globally critical cereal. In Mexico, a significant portion of its annual production is derived from rainfed landraces cultivated by smallholders, but yields are threatened by insect pests, such as the fall armyworm ( Spodoptera frugiperda ). This study evaluated how functional traits of three maize landraces (Nal tel, Nal xoy, Dzit bacal) influence the associated arthropod community and its relationship with S. frugiperda damage. A randomized complete block design was used to assess morphological traits, insect diversity, and foliar damage. The landrace Dzit bacal exhibited superior development in height, stem diameter, leaf area, and leaf dry mass. While no significant differences in pest damage were found among landraces, Nal xoy and Dzit bacal supported arthropod communities with higher ecological diversity (q = 1) and dominance (q = 2), indicating greater evenness. Path analysis identified plant architecture, specifically leaf dry mass, leaf area, and height, as the primary factor structuring the insect community. These results suggest that morphological diversity among landraces acts as a bottom-up driver, shaping arthropod assemblages and favoring natural enemies, such as Ichneumonidae, which may enhance the biological control potential against pests. Fall armyworm leaf damage plant defense insect diversity natural enemies Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Key Message We tested whether maize landrace traits shape insect communities and natural pest control. Plant architecture, not damage levels, is the key driver of structuring arthropod assemblages. Architecturally complex landraces attract natural enemies, such as parasitoid wasps. Maize morphological diversity is a bottom-up tool to enhance biological control resilience. INTRODUCTION Maize ( Zea mays L.) is the most important cereal worldwide. Its annual production reaches 1,027.10 million tons, concentrated in ten leading producers: the United States, China, Brazil, the European Union, Argentina, India, Ukraine, Mexico, South Africa, and Canada, surpassing rice and wheat (Erenstein et al. 2022 ; USDA 2025). In this context, Mexico ranks eighth globally, producing approximately 24 million tons per year. Nearly 58% of this output comes from rainfed systems managed by smallholder farmers who traditionally cultivate landraces adapted to their local conditions (Tanumihardjo et al. 2020 ; SIAP 2025). These landraces are not only central to Mexican gastronomy but also represent a key component of the country’s environmental heritage and rural economy (Palacio-Rojas et al. 2020; Rodríguez-Bustos et al. 2023 ). At the national level, 59 maize races have been documented in Mexico, each of them characterized by broad genetic diversity and adaptability to local agroclimatic conditions and biotic pressures (Hellin et al. 2014 ; Guzzon et al. 2021 ; CONABIO 2025). These landraces are distributed across the entire country, with their highest concentration occurring in the south-central and southern regions, particularly in the states of Oaxaca, Jalisco, Michoacán, and Chiapas (Orozco-Ramírez et al. 2017 ). In the specific case of Yucatán, approximately 5% of the national diversity has been reported, represented mainly by the Tuxpeño , Nal tel , and Dzit bacal landraces, along with their local variants (Arias et al. 2007 ; Ku-Pech et al. 2023 ). Despite its importance to production, maize yield is constrained by damage caused by key insect pests such as the fall armyworm ( Spodoptera frugiperda J.E. Smith), corn leafhopper ( Dalbulus maidis DeLong and Wolcott), corn earworm ( Helicoverpa zea B.), and European corn borer ( Ostrinia nubilalis H.) (Overton et al. 2021 ; Revilla et al. 2021 ). These herbivores affect crop establishment and development, resulting in significant yield losses in maize landraces. The intensity of this damage depends largely on the intrinsic characteristics of each plant, as herbivores respond differentially to the traits expressed by distinct maize landraces; however, these same characteristics may also confer beneficial effects (Stam et al. 2014 ; Costes et al. 2013 ). This relationship between plants and the insects that attack them provides an essential framework for understanding how their interactions are structured and how these processes shape crop dynamics. In this context, the conservation and study of indirect plant defense mechanisms present in maize landraces are crucial not only for ensuring food security, but also for developing more sustainable and resilient integrated management strategies against herbivorous insects (dos Santos et al. 2020 ; Zhou et al. 2024 ). In response to herbivory, plants deploy direct defense mechanisms consisting of morphological structures (such as cuticular waxes, trichomes, and leaf toughness) and secondary metabolites that impair herbivore development (Howe et al. 2008; War et al. 2012 ). Such defenses reduce damage in diverse species, including cotton ( Gossypium hirsutum L.), maize ( Z. mays L.), and African eggplant ( Solanum aethiopicum L .) (Kariyat et al. 2017 ; dos Santos et al. 2020 ; Ali et al. 2021 ). At the same time, plants activate indirect defenses through the emission of herbivore-induced plant volatiles (HIPVs), such as linalool and ꞵ-caryophyllene, which recruit natural enemies (parasitoids and predators) to the site of herbivore feeding (McCormick et al. 2012 ; Ali et al. 2023 ; Wang et al. 2025 ). Several biological control agents are known to respond to these volatiles in maize systems, including Chelonus bifoveolatus S. (Onjura et al. 2025 ), Chelonus insularis C. (Ortiz-Carreon et al. 2019 ), Telenomus podisi A. (Nascimento et al. 2023 ), Cotesia marginiventris C. (Schnee et al. 2006 ), and Campoletis sonorensis C. (de Lange et al. 2016 ). However, the efficiency of this indirect defense depends critically on the availability of shelter and resources, both of which are determined by plant architecture (Costes et al. 2013 ; Hassan et al. 2016 ). Plant functional traits and the associated insect community operate along a cascading pathway: greater structural complexity and vegetation cover enhance the richness of natural enemies and, consequently, suppress herbivore populations (Langellotto and Denno 2004 ; Schlinkert et al. 2015 ; Lucatero et al. 2024 ). This relationship was evidenced by Gontijo et al ( 2010 ), who showed that a greater number of leaves in cucumber ( Cucumis sativus L.) enhanced the activity of the predatory mite Phytoseiulus persimilis A. against the two-spotted spider mite ( Tetranychus urticae K.). This finding suggests that plant structural attributes are a fundamental (and often underestimated) component for optimizing the efficacy of beneficial insects in complex agroecosystems such as those dominated by maize landraces. Nevertheless, despite considerable progress in the study of direct and indirect plant defenses, relatively little is known about how the functional traits present in maize landraces modulate arthropod diversity and, in turn, contribute to reductions in foliar damage (Lucatero et al. 2024 ; Mukanga et al. 2024 ; Waterman et al. 2025 ). Understanding this mechanism is critical because landraces, having evolved under persistent herbivory pressure, may exhibit more robust indirect defense strategies than modern varieties. In this context, this study evaluated how functional traits of three maize landraces (Nal tel, Nal xoy, Dzit bacal) influence the associated arthropod community and its relationship with S. frugiperda damage. MATERIALS AND METHODS Experimental site The experiment was conducted in the Municipality of Conkal, Yucatan (21° 04′ 49.9″ N, 89° 29′ 53.8″ W), from July to October 2023. During this period, the mean monthly temperature was 29.9°C, with a maximum of 36.6°C and a minimum of 23.5°C. Accumulated precipitation was 350 mm. The soil is classified as a Leptosol, containing 0.93% N, and total P, K, Ca, and Mg contents of 2.45, 3.5, 49.38, and 2.63 g kg − 1 , respectively (Ruiz-Santiago et al. 2024 ; dos Santos et al. 2024 ). Plant material Three maize landraces collected from the Xoy region in Peto, Yucatan, were established under field conditions. These landraces are locally known as Nal tel (early-maturing, 65 days to flowering), Nal xoy (intermediate-maturing, 75 days to flowering), and Dzit bacal (late-maturing, 95 days to flowering), and were grown in a monoculture system. The experimental layout followed a randomized complete block design (RCBD) with three replications. Sowing was performed directly by placing two seeds per hill, with a spacing of 0.6 m between plants and 1 m between rows. Each replicate comprised a total of 120 plants per landrace. Experimental plots consisted of three rows, each 12 m in length. Agronomic management included N-P-K fertilization at a total rate of 120-60-00 kg ha − 1 , split into two equal applications administered at 10 and 30 days after emergence (DAE). At 50 DAE, Poliquel® Multi foliar fertilizer was applied at a rate of 2 L ha − 1 . No insecticides were applied to ensure the natural establishment of insect populations on the maize landraces. Plant functional traits and leaf damage in maize landraces Morphological traits of the three maize landraces were evaluated at the flowering stage. A total of 12 plants per landrace were sampled. The measured variables included: plant height (cm), stem diameter (mm), total number of leaves, total leaf area (cm 2 ), leaf dry mass (LDM, g), and specific leaf area (SLA, cm 2 g − 1 ). Plant height was measured from the stem base to the flag leaf using a tape measure (Pretul). Stem diameter was measured at the plant base using a digital caliper (Mitutoyo 500-193-30 CD-12"ASX). Total leaf area was determined via a destructive method, which involved detaching and measuring all leaves using a portable leaf area meter (LI-3000C, LI-COR, Lincoln, NE, USA). The total number of leaves was counted directly on each plant, including senescent (dry) leaves. LDM was obtained by drying samples in an industrial oven at 60°C for 4 days and weighing them on a precision balance (DAMAUS CQT1752GR) until constant weight was reached. Finally, SLA (cm 2 g − 1 ) was calculated as the ratio of total leaf area to LDM. Leaf damage caused by the fall armyworm ( S. frugiperda ) was assessed at 20 and 40 DAE by an experienced evaluator. These time points were selected because they correspond to the crop’s critical vegetative stage (V3-V8), during which whorl infestation has the greatest potential impact (El-Tokhy et al. 2024 ). For this assessment, 36 plants per landrace (12 per replicate) were selected. Damage severity was evaluated on fully expanded young whorl leaves using the 10-point visual scale proposed by Davis et al ( 1992 ), where a score of 0 represents a healthy plant and 9 indicates a severely damaged plant. Insect sampling Sampling was conducted during the advanced growth stage across all maize landraces for nine consecutive days, between 7:00 and 10:00. Mouth aspirators (pooters) were used to sample nine rows (three per replicate), covering a total of 360 individual plants (120 per replicate) across the three landraces. Only insects found resting on plant structures were considered valid samples. Upon collection, insects were transferred to clean vials and preserved in 70% ethanol. Specimens were subsequently classified first to the order level and then to the family level using the general dichotomous keys of Borror y White, (1998). Data analysis Functional trait and leaf damage data were analyzed using Generalized Linear Models (GLMs). The distribution family was selected according to goodness-of-fit criteria, based on the Akaike Information Criterion (AIC) and deviance. A Gamma distribution was used for functional morphological traits, whereas a Poisson distribution was applied for leaf damage; in both cases, a logarithmic link function was specified. A 95% confidence level was adopted, and when significant differences were detected, mean comparisons were carried out using the Bonferroni test (p < 0.05). All statistical analyses were conducted using Infostat version 2020 for Windows. To visualize the structure of the arthropod communities associated with the landraces, rank–abundance curves (Whittaker, 1972 ) were constructed by ordering families from highest to lowest abundance along the x-axis. The extent of this axis reflects the number of families present, with longer axes indicating greater richness. Steeper curves denote assemblages with strong dominance (geometric distributions or log series), whereas gentler slopes (log-normal) indicate higher evenness (Magurran 2004 ). Diversity was then compared among groups using Hill numbers (qD) (Jost 2006 ) for orders q = 0, q = 1, and q = 2. Diversity estimates were calculated with 95% confidence intervals (CI). All diversity analyses were performed using the iNEXT package v.2 (Hsieh et al., 2016 ). Path analysis estimations were conducted using Jamovi v.2.5 to evaluate the complex relationships between cultivars, morphological traits, and leaf damage. Specifically, the model was specified to determine the direct and indirect effects of cultivar characteristics on leaf damage mediated by morphological traits. Model fit was assessed in accordance with the criteria proposed by Hu and Bentler ( 1999 ). We considered the model to have a good fit if the Chi-square test was non-significant (chi 2 , p > 0.05) and the Standardized Root Mean Square Residual (SRMR) was low to 0.08. RESULTS Morphological leaf traits and leaf damage in maize landraces Plant morphological traits showed significant differences (p < 0.05) among the three maize landraces across all evaluated variables, except for the SLA (Fig. 1 ). The Dzit bacal landrace exhibited the highest values for plant height (333.3 cm), stem diameter (27.93 mm), number of leaves (18), total leaf area (11,083.33 cm 2 ), and LDM (76.72 g). In contrast, Nal xoy and Nal tel displayed lower values and reduced vegetative development relative to Dzit bacal . Leaf damage caused by S. frugiperda across the three maize landraces showed no significant differences (p > 0.05) at either evaluation time (Fig. 2 ). At 20 DAE, all landraces ( Nal te l, Nal xoy , and Dzit bacal ) exhibited damage scores ranging from 4.5 to 6.0 on the visual scale (Fig. 2 a). By 40 DAE, damage severity remained within a similar range to that observed in the initial evaluation (4.8–6.0) (Fig. 2 b). Insect community among maize landraces A total of 5,978 individuals were recorded, belonging to 24 families grouped into six orders. Abundance varied notably among landraces: Nal tel hosted the highest number of individuals (n = 3,177; 53.12%), followed by Nal xoy (n = 1,667; 27.88%) and Dzit bacal (n = 1,134; 18.96%). The order Diptera was overwhelmingly dominant, accounting for 94.68% of all captured individuals (n = 5,662), followed by Hemiptera (n = 178; 2.98%) and Hymenoptera (n = 83; 1.39%). The remaining orders (Lepidoptera, Coleoptera, and Dermaptera) jointly represented only 0.92% of the total catch. At the family level, Phoridae (Diptera) dominated the assemblage with 5,231 individuals (87.46% of total abundance), followed by Muscidae (Diptera) with 303 individuals (5.07%) and Pyrrhocoridae (Hemiptera) with 174 individuals (2.91%). The remaining families and their absolute abundances are presented in Table 1 . Diversity indices differed among landraces. Family richness remained similar across landraces (q = 0: Nal tel = 16, Nal xoy = 16, Dzit bacal = 13). However, ecological diversity (q = 1: Nal tel = 1.42, Nal xoy = 2.20, Dzit bacal = 2.27) and the effective number of dominant families (q = 2: Nal tel = 1.14, Nal xoy = 1.47, Dzit bacal = 1.55) revealed clear differences. Specifically, the Dzit bacal and Nal tel landraces differed significantly from Nal xoy based on non-overlapping 95% confidence intervals (Fig. 3 ). Rank–abundance curves revealed a comparable structural pattern across the three communities, characterized by generally low evenness. All landraces exhibited curves with a steep initial slope, indicating strong dominance by a small number of families. In each system ( Nal tel , Nal xoy , and Dzit bacal ), Phoridae was overwhelmingly the most abundant family, followed by Muscidae and Pyrrhocoridae, with a clear separation among their ranked positions. After the fourth or fifth rank, the slope of the curves flattened markedly in all landraces, suggesting that the remaining families (e.g., Formicidae, Noctuidae, Cercopidae, and Ichneumonidae) occurred at similarly low abundances. This pattern reflects greater evenness among the less common taxa within the assemblage. Relationships between morphological traits and insect diversity patterns Plant height showed moderate-to-strong positive correlations with the q = 1 (r = 0.59) and q = 2 (r = 0.65) diversity indices. Its association with species richness (q = 0) was also positive but weak (r = 0.30). A similar pattern emerged for leaf number, which exhibited moderate positive correlations with the diversity of common (q = 1; r = 0.52) and dominant (q = 2; r = 0.54) species, while its relationship with richness (q = 0) remained positive yet weak (r = 0.26). SLA showed no significant linear association with either the q = 1 (r = -0.02) or q = 2 (r = -0.01) indices. Only a weak negative correlation was detected between SLA and total species richness (q = 0; r = -0.29). Finally, leaf damage displayed a divergent pattern. No linear correlations were found with the q = 1 (r = -0.07) or q = 2 (r = 0.02) indices; however, a weak-to-moderate negative correlation emerged between leaf damage and species richness (q = 0; r = -0.35) (Fig. 5 ). Table 1 Taxonomic classification and abundance of insect families sampled in three maize landraces. Order Family Nal tel Nal xoy Dzit bacal Total Diptera Phoridae 2889 1366 976 5231 Muscidae 78 116 109 303 Agromyzidae 29 52 34 115 Otitidae 1 4 5 10 Tabanidae 0 1 1 2 Syrphidae 1 0 0 1 Hemiptera Pyrrhocoridae 37 66 71 174 Cercopidae 0 1 0 1 Cicadellidae 0 0 1 1 Largidae 0 1 0 1 Pentatomidae 0 0 1 1 Hymenoptera Formicidae 27 27 12 66 Ichneumonidae 0 10 6 16 Braconidae 1 0 0 1 Lepidoptera Noctuidae 15 14 9 38 Pyralidae 1 0 0 1 Dermaptera Forficulidae 1 2 1 4 Coleoptera Lycidae 1 3 0 4 Chrysomelidae 1 2 0 3 Anthicidae 0 0 1 1 Carabidae 0 1 0 1 Eucnemidae 1 0 0 1 Nitidulidae 0 1 0 1 Tenebrionidae 1 0 0 1 Total abundance 3084 1667 1227 5978 Table 2 Pearson correlation matrix among plant functional traits, insect diversity metrics (q = 0, q = 1, q = 2), and leaf damage in maize landraces. Variable Height Diameter leaves Leaf area LDM SLA q = 0 q = 1 q = 2 Leaf damage Plant height 1 0.5 0.46 0.78 0.85 -0.14 0.29 0.58 0.64 0.34 Stem diameter 0.5 1 0.33 0.76 0.7 0.14 -0.04 0.43 0.49 0.42 Leaf number 0.46 0.33 1 0.34 0.49 -0.24 0.26 0.51 0.53 0.22 Leaf area 0.78 0.76 0.35 1 0.88 0.25 0.14 0.6 0.67 0.43 LDM 0.85 0.71 0.49 0.88 1 -0.22 0.29 0.65 0.71 0.46 SLA -0.14 0.14 -0.24 0.25 -0.22 1 -0.29 -0.02 -0.01 -0.05 q = 0 0.3 -0.04 0.26 0.14 0.29 -0.29 1 0.67 0.59 -0.35 q = 1 0.59 0.43 0.52 0.61 0.65 -0.02 0.67 1 0.99 -0.07 q = 2 0.65 0.49 0.54 0.67 0.71 -0.01 0.59 0.99 1 0.02 Leaf damage 0.34 0.42 0.22 0.42 0.45 -0.04 -0.35 -0.06 0.01 1 LDM = Leaf Dry Mass; SLA = Specific leaf area; q0 = Richness; q1 = Evenness; q2 = Dominance. The path diagram indicated that Nal xoy and Dzit bacal exhibited consistent associations with functional traits, diversity indices, and leaf damage (Fig. 5 ). Dzit bacal showed negative relationships with LDM (-0.88), total leaf area (-0.97), and plant height (-0.81), while displaying strong positive associations with species richness (q = 0) (1.00) and diversity (q = 1) (1.13). Together with dominance (q = 2) (1.31), these diversity components exerted a direct influence on leaf damage. Similarly, Nal xoy exhibited negative relationships with LDM (-0.75), leaf area (-0.90), and plant height (-0.88), reflecting a parallel structural pattern within the model. DISCUSSION Throughout this study, we evaluated how three maize landraces, differentiated by their functional traits and vegetative architecture, structure arthropod diversity and modulate leaf damage associated with S. frugiperda. Overall, our results indicate that differences in plant height, biomass, and leaf number generate distinct microenvironments (such as canopy shade, humidity, and temperature) that directly influence insect relative abundance, dominance, and taxonomic composition. These structural variations, derived from the distinct developmental cycles of the landraces, suggest that morphological traits act as environmental filters capable of shaping the organization of arthropod communities in maize landraces under the specific agroecological conditions of Southeast Mexico. Morphological leaf traits and leaf damage in maize landraces To understand the ecological consequences of these variations, we analyzed the structural differences among landraces in terms of the functional trade-offs between growth rate and tissue investment. The morphological diversity observed is not incidental; rather, it reflects adaptive strategies tightly aligned with each landrace’s phenology. For example, the late-maturing Dzit bacal invested heavily in vegetative structure (showing the highest values for height, biomass, and leaf area) consistent with a strategy centered on long-term growth. In contrast, Nal tel prioritized rapid development with minimal structural investment, whereas Nal xoy adopted an intermediate strategy (Wright et al. 2004 ). These distinct behaviors embody the classic trade-off between resource conservation and rapid acquisition: plants investing in long-lived, structurally robust tissues experience slower returns but develop denser, more persistent canopies, while fast-growing varieties favor quick leaf turnover at a lower energetic cost (Edwards et al. 2014 ). Such functional divergence extends beyond agronomic relevance, as it directly shapes the physical availability of shelter, the complexity of shaded microhabitats, and the effective foraging surface. Consequently, structurally robust landraces like Dzit bacal do more than accumulate biomass—they engineer a more stable and intricate ecological setting, providing critical resources that differ fundamentally from the open, sparse canopies characteristic of early-maturing varieties (Pérez-Harguindeguy et al. 2016 ). Insect community among maize landraces Given that architectural variation among landraces did not translate into differences in leaf damage, we examined how these structural traits influence the configuration of the arthropod community. Although family richness (q = 0) was similar across landraces, abundance-weighted metrics (q = 1 and q = 2, respectively) showed that Nal xoy and Dzit bacal exhibited higher values, indicating more balanced communities that were less dominated by a few families. The Whittaker (rank-abundance) plots support this interpretation. While Phoridae dominated early in all three landraces, the steeper decline observed in Nal tel indicates stronger dominance, whereas Nal xoy and Dzit bacal displayed a greater relative contribution of secondary families such as Muscidae, Pyrrhocoridae, Formicidae, and Ichneumonidae. The presence of these families is consistent with the structural traits of the plants. Owing to their ecological plasticity, Phoridae benefit from shaded microhabitats and leaf litter within dense canopies (Grundmann et al. 2025 ). Muscidae can exploit moisture, exudates, or organic debris (Cook, 2020 ); Pyrrhocoridae, as herbivorous hemipterans, can feed on plant tissues or seeds (Panizzi and Grazia, 2015 ); and Formicidae act as predators or scavengers, taking advantage of complex microclimates and numerous refuges within a more voluminous canopy (Kuchenbecker et al. 2022 ). However, the most notable difference lies in the exclusive presence of Ichneumonidae in Nal xoy and Dzit bacal , the landraces with the most robust architecture. This hymenopteran family includes parasitoids that require structurally complex microhabitats and stable conditions to locate concealed hosts (such as S. frugiperda) and complete their life cycle (Molina-Ochoa et al. 2003 ). Studies conducted in maize fields in the Yucatán Peninsula have documented that Ichneumonidae communities are influenced by habitat structure, suggesting that plants with greater biomass promote their activity (Orozco-Peón et al. 2019 ). Consequently, these results indicate that although the three landraces share arthropod families, functional architecture modulates the internal distribution of abundances and favors the presence of key natural enemies within the studied assemblage (Gagic et al. 2011 ). The occurrence of Ichneumonidae exclusively in the more complex landraces suggests that these genotypes may sustain more effective top-down biological control mechanisms, consistent with studies showing that crop structure can modulate parasitoid efficacy in maize (Peterson et al. 2016 ). Relationships between morphological traits and insect diversity patterns Our path analysis revealed a clear pattern in which plant architecture functions as the primary axis structuring the arthropod community within the landraces. Among the evaluated morphological traits, LDM, leaf area, and plant height exhibited the strongest correlations with the diversity metrics (qD), as shown in the correlation matrix. These relationships were maintained and further strengthened in the Structural Equation Modeling (SEM), where LDM emerged as the most robust direct predictor of both q = 1 and q = 2, suggesting that greater leaf volume and density increase not only overall abundance but particularly the dominance of certain arthropod groups. This pattern aligns with the notion that greater structural complexity favors dominant groups by increasing the availability of more stable microclimatic gradients (Castagneyrol and Jactel 2012 ; Kambach et al. 2023 ). In contrast, the weak association with q = 0 suggests that richness responds more to specialization filters than to the expansion of physical habitat. Conversely, SLA showed weak or negligible correlations with q = 0, q = 1, and q = 2 in the matrix, consistent with its low influence in the SEM model. This indicates that, at least during these early crop stages, leaf toughness or density does not represent a key functional filter for most recorded families. The absence of clear relationships suggests that leaf quality operates as a secondary axis, likely relevant only to a small subset of species with marked specializations, as proposed by Carmona et al ( 2011 ). A notable aspect is the relationship between the diversity metrics (qD) and leaf damage. Simple correlations show that damage is moderately associated with q = 0, but not with q = 1 or q = 2. However, the SEM detected strong direct effects in opposite directions: q = 2 increases damage, whereas q = 1 reduces it. This divergence between simple correlations and direct effects is statistically plausible and typically arises when relationships are mediated and collinearity exists among structural variables (e.g., LDM, leaf area, plant height), which can obscure direct effects in the correlation matrix (Graham 2003 ; Fan and Chen 2016). However, these interactions represent a specific experimental context and may shift under broader field conditions, where insect behavior is inherently dynamic. Functionally, the positive effect of q = 2 indicates that when the insect community is dominated by several highly abundant families (i.e., a high effective number), herbivore pressure tends to increase, likely due to the accumulation of competing or highly efficient phytophagous guilds (Becerra 2015 ). In contrast, the negative effect of q = 1 suggests that greater evenness (meaning a more homogeneous distribution among the common species) may buffer this damage, probably through higher abundance of natural enemies or moderating interspecific competition (Crowder et al. 2010 ). This pattern suggests that although leaf damage is mediated by indirect pathways originating from plant structural traits, its final expression is modulated by the internal composition of the community, particularly by its degree of dominance and evenness. The exclusive presence of Ichneumonidae in the landraces with higher biomass ( Nal xoy and Dzit bacal ) also supports this hypothesis, as these genotypes create a microhabitat that favors parasitoids, particularly sensitive to vegetation cover and structural complexity (Fornoff et al., 2021 ; Ku-Pech et al. 2023 ). Contrary to our initial hypothesis, we found no direct link between landrace and leaf damage. This suggests that the benefit of architectural complexity is not a simple reduction in herbivory, but rather a restructuring of the arthropod community that may lead to more stable, long-term suppression of pests, potentially manifesting later in the crop cycle or under higher pest pressure. Although no differences in damage levels were detected among landraces at 20 and 40 DAE, these structural conditions may promote more effective biological control at later stages, when host availability increases, as parasitoid frequency and activity shift with the advancing crop cycle and increasing structural complexity (Durocher-Granger et al. 2024 ). Collectively, our results depict a system governed by a dominant structural axis (canopy volume and leaf density) that shapes arthropod abundance and dominance; a secondary leaf-quality axis with limited influence at these early stages; and a leaf-damage response emerging from the interaction between both axes, yet directly modulated by the community’s internal structure (q = 1 and q = 2) once effects are adjusted via SEM. These findings highlight the importance of considering landrace architectural traits as bottom-up management tools and as potential catalysts of top-down processes when aiming to promote structure-sensitive natural enemies, such as ichneumonids. CONCLUSION This study demonstrates that the morphological divergence of maize landraces functions as a key bottom-up force that structures arthropod communities beyond the mere provision of resources. While the greater architectural complexity of late-cycle landraces facilitates the accumulation of dominant groups, our findings show that community evenness (q = 1) and the availability of microhabitats for specific natural enemies (such as Ichneumonidae) are the factors that truly buffer herbivory pressure. Consequently, conserving landraces with contrasting functional strategies (fast-return vs. slow-return) represents not only a safeguard of genetic diversity but also a fundamental ecological strategy to sustain biotic regulation mechanisms and functional resilience within the Milpa system. However, these findings should be interpreted as site-specific insights rather than generalizable behavioral patterns, underscoring the need for research across broader temporal and spatial scales to validate these trends. Declarations Author Contributions : Conceptualization, R.R.R.-S. and J.C.A.-H.; Formal analysis, R.R.R.-S., E.R.-S. and A.D.C.-A.; Methodology, R.R.R.-S., R.D.S.-J. and A.D.C.-A.; Software, R.R.R.-S., Y.M.M.-L and A.D.C.-A.; Supervision, R.R.R.-S and J.C.A.-H.; Writing-original draft, R.R.R.-S.; J.C.A.-H and A.D.C.-A.; Writing-review & editing, R.R.R.-S., J.C.A.-H., E.R.-S., A.L.-P., R.D.S.-J., Y.M.M.-L. and A.D.C.-A. Funding: This research received no external funding. Data Availability Statement: The dataset curated during and/or analyzed in the current study is available from the corresponding author on reasonable request. Acknowledgments: The corresponding author thanks SECIHTI for providing a postdoctoral fellowship to CVU 845968, and the authors thank biologist Lisset Anahí Herrera Poot for her contribution to the fieldwork activities. Conflicts of Interest: The authors declare no conflicts of interest. 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07:56:35","extension":"xml","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":186490,"visible":true,"origin":"","legend":"","description":"","filename":"fa849369f1de4fa89abad8c3efe15b281structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/681ed73e25d3bdb6595dc39d.xml"},{"id":100104843,"identity":"7fa0a58c-fa76-4206-a842-80b90b8f1fac","added_by":"auto","created_at":"2026-01-13 04:37:45","extension":"html","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":200674,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/045c8eff9e272581b25c9080.html"},{"id":100104825,"identity":"f8e2f408-55a2-46de-8020-4eb503e73465","added_by":"auto","created_at":"2026-01-13 04:37:45","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":163861,"visible":true,"origin":"","legend":"\u003cp\u003ePlant morphological traits evaluated at the flowering stage (VT) in three maize landraces. (a) Plant height, (b) stem diameter, (c) total leaf area, (d) leaf number, (e) specific leaf area (SLA) and (f) leaf dry mass (LDM). Different letters above the bars indicate significant differences among landraces (Bonferroni, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05). Df = Degrees of freedom; X\u003csup\u003e2\u003c/sup\u003e = Chi-Square statistic.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/7012e8038a4f803ae8313a16.png"},{"id":100366331,"identity":"0b8df61f-8c96-431f-9156-8a4f5f09e56e","added_by":"auto","created_at":"2026-01-16 07:56:14","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":84393,"visible":true,"origin":"","legend":"\u003cp\u003eLeaf damage caused by fall armyworm (\u003cem\u003eS. frugiperda\u003c/em\u003e) in three maize landraces. (a) Damage observed at 20 DAE and (b) damage observed at 40 DAE. Different letters above the bars indicate significant differences among landraces (Bonferroni, \u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05). Df = Degrees of freedom; X\u003csup\u003e2\u003c/sup\u003e = Chi-Square statistic.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/c29af32e1a6e1fdc76006886.png"},{"id":100104828,"identity":"5a132569-e561-44fc-a6de-e99bb5417529","added_by":"auto","created_at":"2026-01-13 04:37:45","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":141634,"visible":true,"origin":"","legend":"\u003cp\u003eRarefaction curves of effective qD diversity for insect communities in three maize landraces: (a) q0 (richness), (b) q1 (evenness), and (c) q2 (dominance). Shaded areas denote ± 95 % confidence intervals (CI). DB = \u003cem\u003eDzit bacal\u003c/em\u003e; NaT = \u003cem\u003eNal tel\u003c/em\u003e; NaX = \u003cem\u003eNal xoy\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/fe96f968eeecf8de6c173c1a.png"},{"id":100365152,"identity":"aa98eb89-9dc3-4d64-90e6-06e6afb1a933","added_by":"auto","created_at":"2026-01-16 07:54:44","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":171682,"visible":true,"origin":"","legend":"\u003cp\u003eRank-abundance curves of insect families associated with three maize landraces. Data are plotted on a logarithmic scale ordered by family rank. Log10 = base-10 logarithm; ni = abundance of the specific family; N = total abundance of the assemblage.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/f5c044fd0d8f4089edc904a3.png"},{"id":100104831,"identity":"f000ff85-449d-4cf4-82cf-1074110a4188","added_by":"auto","created_at":"2026-01-13 04:37:45","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":266142,"visible":true,"origin":"","legend":"\u003cp\u003ePath analysis of morphological traits and diversity indices on leaf damage in maize landraces. SLA = Specific leaf area; LDM = Leaf Dry Mass; q0 = Richness; q1 = Evenness; q2 = Dominance.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/791b79d6676c8ca367524f23.png"},{"id":100394499,"identity":"088b7443-5411-4a48-bd3d-75fe964346b0","added_by":"auto","created_at":"2026-01-16 11:34:06","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1741661,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8456967/v1/d9f2bb93-c32c-4629-a8d4-31979d044414.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Plant architecture shapes arthropod communities and mediates indirect defense in maize landraces","fulltext":[{"header":"Key Message","content":"\u003cp\u003eWe tested whether maize landrace traits shape insect communities and natural pest control. Plant architecture, not damage levels, is the key driver of structuring arthropod assemblages. Architecturally complex landraces attract natural enemies, such as parasitoid wasps. Maize morphological diversity is a bottom-up tool to enhance biological control resilience.\u003c/p\u003e"},{"header":"INTRODUCTION","content":"\u003cp\u003eMaize (\u003cem\u003eZea mays\u003c/em\u003e L.) is the most important cereal worldwide. Its annual production reaches 1,027.10\u0026nbsp;million tons, concentrated in ten leading producers: the United States, China, Brazil, the European Union, Argentina, India, Ukraine, Mexico, South Africa, and Canada, surpassing rice and wheat (Erenstein et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; USDA 2025). In this context, Mexico ranks eighth globally, producing approximately 24\u0026nbsp;million tons per year. Nearly 58% of this output comes from rainfed systems managed by smallholder farmers who traditionally cultivate landraces adapted to their local conditions (Tanumihardjo et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; SIAP 2025). These landraces are not only central to Mexican gastronomy but also represent a key component of the country\u0026rsquo;s environmental heritage and rural economy (Palacio-Rojas et al. 2020; Rodr\u0026iacute;guez-Bustos et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAt the national level, 59 maize races have been documented in Mexico, each of them characterized by broad genetic diversity and adaptability to local agroclimatic conditions and biotic pressures (Hellin et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Guzzon et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; CONABIO 2025). These landraces are distributed across the entire country, with their highest concentration occurring in the south-central and southern regions, particularly in the states of Oaxaca, Jalisco, Michoac\u0026aacute;n, and Chiapas (Orozco-Ram\u0026iacute;rez et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). In the specific case of Yucat\u0026aacute;n, approximately 5% of the national diversity has been reported, represented mainly by the \u003cem\u003eTuxpe\u0026ntilde;o\u003c/em\u003e, \u003cem\u003eNal tel\u003c/em\u003e, and \u003cem\u003eDzit bacal\u003c/em\u003e landraces, along with their local variants (Arias et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Ku-Pech et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eDespite its importance to production, maize yield is constrained by damage caused by key insect pests such as the fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e J.E. Smith), corn leafhopper (\u003cem\u003eDalbulus maidis\u003c/em\u003e DeLong and Wolcott), corn earworm (\u003cem\u003eHelicoverpa zea\u003c/em\u003e B.), and European corn borer (\u003cem\u003eOstrinia nubilalis\u003c/em\u003e H.) (Overton et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Revilla et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These herbivores affect crop establishment and development, resulting in significant yield losses in maize landraces. The intensity of this damage depends largely on the intrinsic characteristics of each plant, as herbivores respond differentially to the traits expressed by distinct maize landraces; however, these same characteristics may also confer beneficial effects (Stam et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Costes et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThis relationship between plants and the insects that attack them provides an essential framework for understanding how their interactions are structured and how these processes shape crop dynamics. In this context, the conservation and study of indirect plant defense mechanisms present in maize landraces are crucial not only for ensuring food security, but also for developing more sustainable and resilient integrated management strategies against herbivorous insects (dos Santos et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Zhou et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn response to herbivory, plants deploy direct defense mechanisms consisting of morphological structures (such as cuticular waxes, trichomes, and leaf toughness) and secondary metabolites that impair herbivore development (Howe et al. 2008; War et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Such defenses reduce damage in diverse species, including cotton (\u003cem\u003eGossypium hirsutum\u003c/em\u003e L.), maize (\u003cem\u003eZ. mays\u003c/em\u003e L.), and African eggplant (\u003cem\u003eSolanum aethiopicum L\u003c/em\u003e.) (Kariyat et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; dos Santos et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ali et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). At the same time, plants activate indirect defenses through the emission of herbivore-induced plant volatiles (HIPVs), such as linalool and ꞵ-caryophyllene, which recruit natural enemies (parasitoids and predators) to the site of herbivore feeding (McCormick et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Ali et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral biological control agents are known to respond to these volatiles in maize systems, including \u003cem\u003eChelonus bifoveolatus\u003c/em\u003e S. (Onjura et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), \u003cem\u003eChelonus insularis\u003c/em\u003e C. (Ortiz-Carreon et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), \u003cem\u003eTelenomus podisi\u003c/em\u003e A. (Nascimento et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), \u003cem\u003eCotesia marginiventris\u003c/em\u003e C. (Schnee et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), and \u003cem\u003eCampoletis sonorensis\u003c/em\u003e C. (de Lange et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). However, the efficiency of this indirect defense depends critically on the availability of shelter and resources, both of which are determined by plant architecture (Costes et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hassan et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Plant functional traits and the associated insect community operate along a cascading pathway: greater structural complexity and vegetation cover enhance the richness of natural enemies and, consequently, suppress herbivore populations (Langellotto and Denno \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Schlinkert et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Lucatero et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThis relationship was evidenced by Gontijo et al (\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), who showed that a greater number of leaves in cucumber (\u003cem\u003eCucumis sativus\u003c/em\u003e L.) enhanced the activity of the predatory mite \u003cem\u003ePhytoseiulus persimilis\u003c/em\u003e A. against the two-spotted spider mite (\u003cem\u003eTetranychus urticae\u003c/em\u003e K.). This finding suggests that plant structural attributes are a fundamental (and often underestimated) component for optimizing the efficacy of beneficial insects in complex agroecosystems such as those dominated by maize landraces. Nevertheless, despite considerable progress in the study of direct and indirect plant defenses, relatively little is known about how the functional traits present in maize landraces modulate arthropod diversity and, in turn, contribute to reductions in foliar damage (Lucatero et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Mukanga et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Waterman et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Understanding this mechanism is critical because landraces, having evolved under persistent herbivory pressure, may exhibit more robust indirect defense strategies than modern varieties. In this context, this study evaluated how functional traits of three maize landraces (Nal tel, Nal xoy, Dzit bacal) influence the associated arthropod community and its relationship with \u003cem\u003eS. frugiperda\u003c/em\u003e damage.\u003c/p\u003e"},{"header":"MATERIALS AND METHODS","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eExperimental site\u003c/h2\u003e \u003cp\u003eThe experiment was conducted in the Municipality of Conkal, Yucatan (21\u0026deg; 04\u0026prime; 49.9\u0026Prime; N, 89\u0026deg; 29\u0026prime; 53.8\u0026Prime; W), from July to October 2023. During this period, the mean monthly temperature was 29.9\u0026deg;C, with a maximum of 36.6\u0026deg;C and a minimum of 23.5\u0026deg;C. Accumulated precipitation was 350 mm. The soil is classified as a Leptosol, containing 0.93% N, and total P, K, Ca, and Mg contents of 2.45, 3.5, 49.38, and 2.63 g kg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, respectively (Ruiz-Santiago et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; dos Santos et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003ePlant material\u003c/h3\u003e\n\u003cp\u003eThree maize landraces collected from the Xoy region in Peto, Yucatan, were established under field conditions. These landraces are locally known as \u003cem\u003eNal tel\u003c/em\u003e (early-maturing, 65 days to flowering), \u003cem\u003eNal xoy\u003c/em\u003e (intermediate-maturing, 75 days to flowering), and \u003cem\u003eDzit bacal\u003c/em\u003e (late-maturing, 95 days to flowering), and were grown in a monoculture system. The experimental layout followed a randomized complete block design (RCBD) with three replications.\u003c/p\u003e \u003cp\u003eSowing was performed directly by placing two seeds per hill, with a spacing of 0.6 m between plants and 1 m between rows. Each replicate comprised a total of 120 plants per landrace. Experimental plots consisted of three rows, each 12 m in length. Agronomic management included N-P-K fertilization at a total rate of 120-60-00 kg ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, split into two equal applications administered at 10 and 30 days after emergence (DAE). At 50 DAE, Poliquel\u0026reg; Multi foliar fertilizer was applied at a rate of 2 L ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. No insecticides were applied to ensure the natural establishment of insect populations on the maize landraces.\u003c/p\u003e\n\u003ch3\u003ePlant functional traits and leaf damage in maize landraces\u003c/h3\u003e\n\u003cp\u003eMorphological traits of the three maize landraces were evaluated at the flowering stage. A total of 12 plants per landrace were sampled. The measured variables included: plant height (cm), stem diameter (mm), total number of leaves, total leaf area (cm\u003csup\u003e2\u003c/sup\u003e), leaf dry mass (LDM, g), and specific leaf area (SLA, cm\u003csup\u003e2\u003c/sup\u003e g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e). Plant height was measured from the stem base to the flag leaf using a tape measure (Pretul). Stem diameter was measured at the plant base using a digital caliper (Mitutoyo 500-193-30 CD-12\"ASX). Total leaf area was determined via a destructive method, which involved detaching and measuring all leaves using a portable leaf area meter (LI-3000C, LI-COR, Lincoln, NE, USA). The total number of leaves was counted directly on each plant, including senescent (dry) leaves. LDM was obtained by drying samples in an industrial oven at 60\u0026deg;C for 4 days and weighing them on a precision balance (DAMAUS CQT1752GR) until constant weight was reached. Finally, SLA (cm\u003csup\u003e2\u003c/sup\u003e g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) was calculated as the ratio of total leaf area to LDM.\u003c/p\u003e \u003cp\u003eLeaf damage caused by the fall armyworm (\u003cem\u003eS. frugiperda\u003c/em\u003e) was assessed at 20 and 40 DAE by an experienced evaluator. These time points were selected because they correspond to the crop\u0026rsquo;s critical vegetative stage (V3-V8), during which whorl infestation has the greatest potential impact (El-Tokhy et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). For this assessment, 36 plants per landrace (12 per replicate) were selected. Damage severity was evaluated on fully expanded young whorl leaves using the 10-point visual scale proposed by Davis et al (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1992\u003c/span\u003e), where a score of 0 represents a healthy plant and 9 indicates a severely damaged plant.\u003c/p\u003e\n\u003ch3\u003eInsect sampling\u003c/h3\u003e\n\u003cp\u003eSampling was conducted during the advanced growth stage across all maize landraces for nine consecutive days, between 7:00 and 10:00. Mouth aspirators (pooters) were used to sample nine rows (three per replicate), covering a total of 360 individual plants (120 per replicate) across the three landraces. Only insects found resting on plant structures were considered valid samples.\u003c/p\u003e \u003cp\u003eUpon collection, insects were transferred to clean vials and preserved in 70% ethanol. Specimens were subsequently classified first to the order level and then to the family level using the general dichotomous keys of Borror y White, (1998).\u003c/p\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eFunctional trait and leaf damage data were analyzed using Generalized Linear Models (GLMs). The distribution family was selected according to goodness-of-fit criteria, based on the Akaike Information Criterion (AIC) and deviance. A Gamma distribution was used for functional morphological traits, whereas a Poisson distribution was applied for leaf damage; in both cases, a logarithmic link function was specified. A 95% confidence level was adopted, and when significant differences were detected, mean comparisons were carried out using the Bonferroni test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). All statistical analyses were conducted using Infostat version 2020 for Windows.\u003c/p\u003e \u003cp\u003eTo visualize the structure of the arthropod communities associated with the landraces, rank\u0026ndash;abundance curves (Whittaker, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1972\u003c/span\u003e) were constructed by ordering families from highest to lowest abundance along the x-axis. The extent of this axis reflects the number of families present, with longer axes indicating greater richness. Steeper curves denote assemblages with strong dominance (geometric distributions or log series), whereas gentler slopes (log-normal) indicate higher evenness (Magurran \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Diversity was then compared among groups using Hill numbers (qD) (Jost \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) for orders q\u0026thinsp;=\u0026thinsp;0, q\u0026thinsp;=\u0026thinsp;1, and q\u0026thinsp;=\u0026thinsp;2. Diversity estimates were calculated with 95% confidence intervals (CI). All diversity analyses were performed using the iNEXT package v.2 (Hsieh et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePath analysis estimations were conducted using Jamovi v.2.5 to evaluate the complex relationships between cultivars, morphological traits, and leaf damage. Specifically, the model was specified to determine the direct and indirect effects of cultivar characteristics on leaf damage mediated by morphological traits. Model fit was assessed in accordance with the criteria proposed by Hu and Bentler (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). We considered the model to have a good fit if the Chi-square test was non-significant (chi\u003csup\u003e2\u003c/sup\u003e, p\u0026thinsp;\u0026gt;\u0026thinsp;0.05) and the Standardized Root Mean Square Residual (SRMR) was low to 0.08.\u003c/p\u003e \u003c/div\u003e"},{"header":"RESULTS","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eMorphological leaf traits and leaf damage in maize landraces\u003c/h2\u003e \u003cp\u003ePlant morphological traits showed significant differences (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) among the three maize landraces across all evaluated variables, except for the SLA (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The \u003cem\u003eDzit bacal\u003c/em\u003e landrace exhibited the highest values for plant height (333.3 cm), stem diameter (27.93 mm), number of leaves (18), total leaf area (11,083.33 cm\u003csup\u003e2\u003c/sup\u003e), and LDM (76.72 g). In contrast, \u003cem\u003eNal xoy\u003c/em\u003e and \u003cem\u003eNal tel\u003c/em\u003e displayed lower values and reduced vegetative development relative to \u003cem\u003eDzit bacal\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eLeaf damage caused by \u003cem\u003eS. frugiperda\u003c/em\u003e across the three maize landraces showed no significant differences (p\u0026thinsp;\u0026gt;\u0026thinsp;0.05) at either evaluation time (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). At 20 DAE, all landraces (\u003cem\u003eNal te\u003c/em\u003el, \u003cem\u003eNal xoy\u003c/em\u003e, and \u003cem\u003eDzit bacal\u003c/em\u003e) exhibited damage scores ranging from 4.5 to 6.0 on the visual scale (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). By 40 DAE, damage severity remained within a similar range to that observed in the initial evaluation (4.8\u0026ndash;6.0) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eInsect community among maize landraces\u003c/h3\u003e\n\u003cp\u003eA total of 5,978 individuals were recorded, belonging to 24 families grouped into six orders. Abundance varied notably among landraces: \u003cem\u003eNal tel\u003c/em\u003e hosted the highest number of individuals (n\u0026thinsp;=\u0026thinsp;3,177; 53.12%), followed by \u003cem\u003eNal xoy\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;1,667; 27.88%) and \u003cem\u003eDzit bacal\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;1,134; 18.96%). The order Diptera was overwhelmingly dominant, accounting for 94.68% of all captured individuals (n\u0026thinsp;=\u0026thinsp;5,662), followed by Hemiptera (n\u0026thinsp;=\u0026thinsp;178; 2.98%) and Hymenoptera (n\u0026thinsp;=\u0026thinsp;83; 1.39%). The remaining orders (Lepidoptera, Coleoptera, and Dermaptera) jointly represented only 0.92% of the total catch. At the family level, Phoridae (Diptera) dominated the assemblage with 5,231 individuals (87.46% of total abundance), followed by Muscidae (Diptera) with 303 individuals (5.07%) and Pyrrhocoridae (Hemiptera) with 174 individuals (2.91%). The remaining families and their absolute abundances are presented in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Diversity indices differed among landraces. Family richness remained similar across landraces (q\u0026thinsp;=\u0026thinsp;0: \u003cem\u003eNal tel\u003c/em\u003e\u0026thinsp;=\u0026thinsp;16, \u003cem\u003eNal xoy\u003c/em\u003e\u0026thinsp;=\u0026thinsp;16, \u003cem\u003eDzit bacal\u003c/em\u003e\u0026thinsp;=\u0026thinsp;13). However, ecological diversity (q\u0026thinsp;=\u0026thinsp;1: \u003cem\u003eNal tel\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.42, \u003cem\u003eNal xoy\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.20, \u003cem\u003eDzit bacal\u003c/em\u003e\u0026thinsp;=\u0026thinsp;2.27) and the effective number of dominant families (q\u0026thinsp;=\u0026thinsp;2: \u003cem\u003eNal tel\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.14, \u003cem\u003eNal xoy\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.47, \u003cem\u003eDzit bacal\u003c/em\u003e\u0026thinsp;=\u0026thinsp;1.55) revealed clear differences. Specifically, the \u003cem\u003eDzit bacal\u003c/em\u003e and \u003cem\u003eNal tel\u003c/em\u003e landraces differed significantly from \u003cem\u003eNal xoy\u003c/em\u003e based on non-overlapping 95% confidence intervals (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRank\u0026ndash;abundance curves revealed a comparable structural pattern across the three communities, characterized by generally low evenness. All landraces exhibited curves with a steep initial slope, indicating strong dominance by a small number of families. In each system (\u003cem\u003eNal tel\u003c/em\u003e, \u003cem\u003eNal xoy\u003c/em\u003e, and \u003cem\u003eDzit bacal\u003c/em\u003e), Phoridae was overwhelmingly the most abundant family, followed by Muscidae and Pyrrhocoridae, with a clear separation among their ranked positions. After the fourth or fifth rank, the slope of the curves flattened markedly in all landraces, suggesting that the remaining families (e.g., Formicidae, Noctuidae, Cercopidae, and Ichneumonidae) occurred at similarly low abundances. This pattern reflects greater evenness among the less common taxa within the assemblage.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eRelationships between morphological traits and insect diversity patterns\u003c/h2\u003e \u003cp\u003ePlant height showed moderate-to-strong positive correlations with the q\u0026thinsp;=\u0026thinsp;1 (r\u0026thinsp;=\u0026thinsp;0.59) and q\u0026thinsp;=\u0026thinsp;2 (r\u0026thinsp;=\u0026thinsp;0.65) diversity indices. Its association with species richness (q\u0026thinsp;=\u0026thinsp;0) was also positive but weak (r\u0026thinsp;=\u0026thinsp;0.30). A similar pattern emerged for leaf number, which exhibited moderate positive correlations with the diversity of common (q\u0026thinsp;=\u0026thinsp;1; r\u0026thinsp;=\u0026thinsp;0.52) and dominant (q\u0026thinsp;=\u0026thinsp;2; r\u0026thinsp;=\u0026thinsp;0.54) species, while its relationship with richness (q\u0026thinsp;=\u0026thinsp;0) remained positive yet weak (r\u0026thinsp;=\u0026thinsp;0.26).\u003c/p\u003e \u003cp\u003eSLA showed no significant linear association with either the q\u0026thinsp;=\u0026thinsp;1 (r = -0.02) or q\u0026thinsp;=\u0026thinsp;2 (r = -0.01) indices. Only a weak negative correlation was detected between SLA and total species richness (q\u0026thinsp;=\u0026thinsp;0; r = -0.29). Finally, leaf damage displayed a divergent pattern. No linear correlations were found with the q\u0026thinsp;=\u0026thinsp;1 (r = -0.07) or q\u0026thinsp;=\u0026thinsp;2 (r\u0026thinsp;=\u0026thinsp;0.02) indices; however, a weak-to-moderate negative correlation emerged between leaf damage and species richness (q\u0026thinsp;=\u0026thinsp;0; r = -0.35) (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\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\u003eTaxonomic classification and abundance of insect families sampled in three maize landraces.\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=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOrder\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFamily\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003eNal tel\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eNal xoy\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eDzit bacal\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDiptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePhoridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2889\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1366\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e976\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e5231\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMuscidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e303\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAgromyzidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e115\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eOtitidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTabanidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSyrphidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHemiptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePyrrhocoridae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e174\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCercopidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCicadellidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLargidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePentatomidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHymenoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFormicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eIchneumonidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBraconidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLepidoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNoctuidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePyralidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDermaptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eForficulidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eColeoptera\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLycidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eChrysomelidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAnthicidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCarabidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEucnemidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNitidulidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTenebrionidae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal abundance\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e3084\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e1667\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e1227\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e5978\u003c/b\u003e\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 \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePearson correlation matrix among plant functional traits, insect diversity metrics (q\u0026thinsp;=\u0026thinsp;0, q\u0026thinsp;=\u0026thinsp;1, q\u0026thinsp;=\u0026thinsp;2), and leaf damage in maize landraces.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVariable\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHeight\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eDiameter\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eleaves\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLeaf area\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLDM\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSLA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eq\u0026thinsp;=\u0026thinsp;0\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eq\u0026thinsp;=\u0026thinsp;1\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eq\u0026thinsp;=\u0026thinsp;2\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eLeaf damage\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePlant height\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.78\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.85\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.58\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.64\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eStem diameter\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.76\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.42\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLeaf number\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.51\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.53\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.22\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLeaf area\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e0.78\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e0.76\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.88\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.67\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.43\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLDM\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e0.85\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e0.71\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.88\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.71\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.46\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSLA\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e-0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e-0.05\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eq\u0026thinsp;=\u0026thinsp;0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e-0.35\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eq\u0026thinsp;=\u0026thinsp;1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e0.59\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e0.52\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.61\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e0.99\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e-0.07\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eq\u0026thinsp;=\u0026thinsp;2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e0.65\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e0.54\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e0.67\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e0.71\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.99\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.02\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eLeaf damage\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\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\u003eLDM\u0026thinsp;=\u0026thinsp;Leaf Dry Mass; SLA\u0026thinsp;=\u0026thinsp;Specific leaf area; q0\u0026thinsp;=\u0026thinsp;Richness; q1\u0026thinsp;=\u0026thinsp;Evenness; q2\u0026thinsp;=\u0026thinsp;Dominance.\u003c/p\u003e \u003cp\u003eThe path diagram indicated that \u003cem\u003eNal xoy\u003c/em\u003e and \u003cem\u003eDzit bacal\u003c/em\u003e exhibited consistent associations with functional traits, diversity indices, and leaf damage (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). \u003cem\u003eDzit bacal\u003c/em\u003e showed negative relationships with LDM (-0.88), total leaf area (-0.97), and plant height (-0.81), while displaying strong positive associations with species richness (q\u0026thinsp;=\u0026thinsp;0) (1.00) and diversity (q\u0026thinsp;=\u0026thinsp;1) (1.13). Together with dominance (q\u0026thinsp;=\u0026thinsp;2) (1.31), these diversity components exerted a direct influence on leaf damage. Similarly, \u003cem\u003eNal xoy\u003c/em\u003e exhibited negative relationships with LDM (-0.75), leaf area (-0.90), and plant height (-0.88), reflecting a parallel structural pattern within the model.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eThroughout this study, we evaluated how three maize landraces, differentiated by their functional traits and vegetative architecture, structure arthropod diversity and modulate leaf damage associated with S. frugiperda. Overall, our results indicate that differences in plant height, biomass, and leaf number generate distinct microenvironments (such as canopy shade, humidity, and temperature) that directly influence insect relative abundance, dominance, and taxonomic composition. These structural variations, derived from the distinct developmental cycles of the landraces, suggest that morphological traits act as environmental filters capable of shaping the organization of arthropod communities in maize landraces under the specific agroecological conditions of Southeast Mexico.\u003c/p\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eMorphological leaf traits and leaf damage in maize landraces\u003c/h2\u003e \u003cp\u003eTo understand the ecological consequences of these variations, we analyzed the structural differences among landraces in terms of the functional trade-offs between growth rate and tissue investment. The morphological diversity observed is not incidental; rather, it reflects adaptive strategies tightly aligned with each landrace\u0026rsquo;s phenology. For example, the late-maturing \u003cem\u003eDzit bacal\u003c/em\u003e invested heavily in vegetative structure (showing the highest values for height, biomass, and leaf area) consistent with a strategy centered on long-term growth. In contrast, \u003cem\u003eNal tel\u003c/em\u003e prioritized rapid development with minimal structural investment, whereas \u003cem\u003eNal xoy\u003c/em\u003e adopted an intermediate strategy (Wright et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). These distinct behaviors embody the classic trade-off between resource conservation and rapid acquisition: plants investing in long-lived, structurally robust tissues experience slower returns but develop denser, more persistent canopies, while fast-growing varieties favor quick leaf turnover at a lower energetic cost (Edwards et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSuch functional divergence extends beyond agronomic relevance, as it directly shapes the physical availability of shelter, the complexity of shaded microhabitats, and the effective foraging surface. Consequently, structurally robust landraces like Dzit bacal do more than accumulate biomass\u0026mdash;they engineer a more stable and intricate ecological setting, providing critical resources that differ fundamentally from the open, sparse canopies characteristic of early-maturing varieties (P\u0026eacute;rez-Harguindeguy et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eInsect community among maize landraces\u003c/h2\u003e \u003cp\u003eGiven that architectural variation among landraces did not translate into differences in leaf damage, we examined how these structural traits influence the configuration of the arthropod community. Although family richness (q\u0026thinsp;=\u0026thinsp;0) was similar across landraces, abundance-weighted metrics (q\u0026thinsp;=\u0026thinsp;1 and q\u0026thinsp;=\u0026thinsp;2, respectively) showed that \u003cem\u003eNal xoy\u003c/em\u003e and \u003cem\u003eDzit bacal\u003c/em\u003e exhibited higher values, indicating more balanced communities that were less dominated by a few families.\u003c/p\u003e \u003cp\u003eThe Whittaker (rank-abundance) plots support this interpretation. While Phoridae dominated early in all three landraces, the steeper decline observed in \u003cem\u003eNal tel\u003c/em\u003e indicates stronger dominance, whereas \u003cem\u003eNal xoy\u003c/em\u003e and \u003cem\u003eDzit bacal\u003c/em\u003e displayed a greater relative contribution of secondary families such as Muscidae, Pyrrhocoridae, Formicidae, and Ichneumonidae.\u003c/p\u003e \u003cp\u003eThe presence of these families is consistent with the structural traits of the plants. Owing to their ecological plasticity, Phoridae benefit from shaded microhabitats and leaf litter within dense canopies (Grundmann et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Muscidae can exploit moisture, exudates, or organic debris (Cook, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e); Pyrrhocoridae, as herbivorous hemipterans, can feed on plant tissues or seeds (Panizzi and Grazia, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2015\u003c/span\u003e); and Formicidae act as predators or scavengers, taking advantage of complex microclimates and numerous refuges within a more voluminous canopy (Kuchenbecker et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHowever, the most notable difference lies in the exclusive presence of Ichneumonidae in \u003cem\u003eNal xoy\u003c/em\u003e and \u003cem\u003eDzit bacal\u003c/em\u003e, the landraces with the most robust architecture. This hymenopteran family includes parasitoids that require structurally complex microhabitats and stable conditions to locate concealed hosts (such as S. frugiperda) and complete their life cycle (Molina-Ochoa et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Studies conducted in maize fields in the Yucat\u0026aacute;n Peninsula have documented that Ichneumonidae communities are influenced by habitat structure, suggesting that plants with greater biomass promote their activity (Orozco-Pe\u0026oacute;n et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eConsequently, these results indicate that although the three landraces share arthropod families, functional architecture modulates the internal distribution of abundances and favors the presence of key natural enemies within the studied assemblage (Gagic et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). The occurrence of Ichneumonidae exclusively in the more complex landraces suggests that these genotypes may sustain more effective top-down biological control mechanisms, consistent with studies showing that crop structure can modulate parasitoid efficacy in maize (Peterson et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eRelationships between morphological traits and insect diversity patterns\u003c/h2\u003e \u003cp\u003eOur path analysis revealed a clear pattern in which plant architecture functions as the primary axis structuring the arthropod community within the landraces. Among the evaluated morphological traits, LDM, leaf area, and plant height exhibited the strongest correlations with the diversity metrics (qD), as shown in the correlation matrix. These relationships were maintained and further strengthened in the Structural Equation Modeling (SEM), where LDM emerged as the most robust direct predictor of both q\u0026thinsp;=\u0026thinsp;1 and q\u0026thinsp;=\u0026thinsp;2, suggesting that greater leaf volume and density increase not only overall abundance but particularly the dominance of certain arthropod groups. This pattern aligns with the notion that greater structural complexity favors dominant groups by increasing the availability of more stable microclimatic gradients (Castagneyrol and Jactel \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Kambach et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In contrast, the weak association with q\u0026thinsp;=\u0026thinsp;0 suggests that richness responds more to specialization filters than to the expansion of physical habitat.\u003c/p\u003e \u003cp\u003eConversely, SLA showed weak or negligible correlations with q\u0026thinsp;=\u0026thinsp;0, q\u0026thinsp;=\u0026thinsp;1, and q\u0026thinsp;=\u0026thinsp;2 in the matrix, consistent with its low influence in the SEM model. This indicates that, at least during these early crop stages, leaf toughness or density does not represent a key functional filter for most recorded families. The absence of clear relationships suggests that leaf quality operates as a secondary axis, likely relevant only to a small subset of species with marked specializations, as proposed by Carmona et al (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA notable aspect is the relationship between the diversity metrics (qD) and leaf damage. Simple correlations show that damage is moderately associated with q\u0026thinsp;=\u0026thinsp;0, but not with q\u0026thinsp;=\u0026thinsp;1 or q\u0026thinsp;=\u0026thinsp;2. However, the SEM detected strong direct effects in opposite directions: q\u0026thinsp;=\u0026thinsp;2 increases damage, whereas q\u0026thinsp;=\u0026thinsp;1 reduces it. This divergence between simple correlations and direct effects is statistically plausible and typically arises when relationships are mediated and collinearity exists among structural variables (e.g., LDM, leaf area, plant height), which can obscure direct effects in the correlation matrix (Graham \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Fan and Chen 2016). However, these interactions represent a specific experimental context and may shift under broader field conditions, where insect behavior is inherently dynamic.\u003c/p\u003e \u003cp\u003eFunctionally, the positive effect of q\u0026thinsp;=\u0026thinsp;2 indicates that when the insect community is dominated by several highly abundant families (i.e., a high effective number), herbivore pressure tends to increase, likely due to the accumulation of competing or highly efficient phytophagous guilds (Becerra \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). In contrast, the negative effect of q\u0026thinsp;=\u0026thinsp;1 suggests that greater evenness (meaning a more homogeneous distribution among the common species) may buffer this damage, probably through higher abundance of natural enemies or moderating interspecific competition (Crowder et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). This pattern suggests that although leaf damage is mediated by indirect pathways originating from plant structural traits, its final expression is modulated by the internal composition of the community, particularly by its degree of dominance and evenness.\u003c/p\u003e \u003cp\u003eThe exclusive presence of Ichneumonidae in the landraces with higher biomass (\u003cem\u003eNal xoy\u003c/em\u003e and \u003cem\u003eDzit bacal\u003c/em\u003e) also supports this hypothesis, as these genotypes create a microhabitat that favors parasitoids, particularly sensitive to vegetation cover and structural complexity (Fornoff et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Ku-Pech et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Contrary to our initial hypothesis, we found no direct link between landrace and leaf damage. This suggests that the benefit of architectural complexity is not a simple reduction in herbivory, but rather a restructuring of the arthropod community that may lead to more stable, long-term suppression of pests, potentially manifesting later in the crop cycle or under higher pest pressure. Although no differences in damage levels were detected among landraces at 20 and 40 DAE, these structural conditions may promote more effective biological control at later stages, when host availability increases, as parasitoid frequency and activity shift with the advancing crop cycle and increasing structural complexity (Durocher-Granger et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCollectively, our results depict a system governed by a dominant structural axis (canopy volume and leaf density) that shapes arthropod abundance and dominance; a secondary leaf-quality axis with limited influence at these early stages; and a leaf-damage response emerging from the interaction between both axes, yet directly modulated by the community\u0026rsquo;s internal structure (q\u0026thinsp;=\u0026thinsp;1 and q\u0026thinsp;=\u0026thinsp;2) once effects are adjusted via SEM. These findings highlight the importance of considering landrace architectural traits as bottom-up management tools and as potential catalysts of top-down processes when aiming to promote structure-sensitive natural enemies, such as ichneumonids.\u003c/p\u003e \u003c/div\u003e"},{"header":"CONCLUSION","content":"\u003cp\u003eThis study demonstrates that the morphological divergence of maize landraces functions as a key bottom-up force that structures arthropod communities beyond the mere provision of resources. While the greater architectural complexity of late-cycle landraces facilitates the accumulation of dominant groups, our findings show that community evenness (q\u0026thinsp;=\u0026thinsp;1) and the availability of microhabitats for specific natural enemies (such as Ichneumonidae) are the factors that truly buffer herbivory pressure. Consequently, conserving landraces with contrasting functional strategies (fast-return vs. slow-return) represents not only a safeguard of genetic diversity but also a fundamental ecological strategy to sustain biotic regulation mechanisms and functional resilience within the Milpa system. However, these findings should be interpreted as site-specific insights rather than generalizable behavioral patterns, underscoring the need for research across broader temporal and spatial scales to validate these trends.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e: Conceptualization, R.R.R.-S. and J.C.A.-H.; Formal analysis, R.R.R.-S., E.R.-S. and A.D.C.-A.; Methodology, R.R.R.-S., R.D.S.-J. and A.D.C.-A.; Software, R.R.R.-S., Y.M.M.-L and A.D.C.-A.; Supervision, R.R.R.-S and J.C.A.-H.; Writing-original draft, R.R.R.-S.; J.C.A.-H and A.D.C.-A.; Writing-review \u0026amp; editing, R.R.R.-S., J.C.A.-H., E.R.-S., A.L.-P., R.D.S.-J., Y.M.M.-L. and A.D.C.-A.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u003c/strong\u003e This research received no external funding.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability Statement:\u003c/strong\u003e The dataset curated during and/or analyzed in the current study is available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgments:\u0026nbsp;\u003c/strong\u003eThe corresponding author thanks SECIHTI for providing a postdoctoral fellowship to CVU 845968, and the authors thank biologist Lisset Anah\u0026iacute; Herrera Poot for her contribution to the fieldwork activities.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of Interest:\u003c/strong\u003e The authors declare no conflicts of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAli MA, Khan MAU, Rao AQ, Iqbal A, Din SU, Shahid AA (2021) Biochemical evidence of epicuticular wax compounds involved in cotton-whitefly interaction. 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ACS Omega 9(40): 41130-41147. https://doi.org/10.1021/acsomega.4c05847\u003c/li\u003e\n\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":"Fall armyworm, leaf damage, plant defense, insect diversity, natural enemies","lastPublishedDoi":"10.21203/rs.3.rs-8456967/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8456967/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMaize (\u003cem\u003eZea mays\u003c/em\u003e L.) is a globally critical cereal. In Mexico, a significant portion of its annual production is derived from rainfed landraces cultivated by smallholders, but yields are threatened by insect pests, such as the fall armyworm (\u003cem\u003eSpodoptera frugiperda\u003c/em\u003e). This study evaluated how functional traits of three maize landraces (Nal tel, Nal xoy, Dzit bacal) influence the associated arthropod community and its relationship with \u003cem\u003eS. frugiperda\u003c/em\u003e damage. A randomized complete block design was used to assess morphological traits, insect diversity, and foliar damage. The landrace Dzit bacal exhibited superior development in height, stem diameter, leaf area, and leaf dry mass. While no significant differences in pest damage were found among landraces, Nal xoy and Dzit bacal supported arthropod communities with higher ecological diversity (q = 1) and dominance (q = 2), indicating greater evenness. Path analysis identified plant architecture, specifically leaf dry mass, leaf area, and height, as the primary factor structuring the insect community. These results suggest that morphological diversity among landraces acts as a bottom-up driver, shaping arthropod assemblages and favoring natural enemies, such as Ichneumonidae, which may enhance the biological control potential against pests.\u003c/p\u003e","manuscriptTitle":"Plant architecture shapes arthropod communities and mediates indirect defense in maize landraces","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-01-13 04:37:40","doi":"10.21203/rs.3.rs-8456967/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":"2814052f-e0d1-4906-b21a-7e94094d20e9","owner":[],"postedDate":"January 13th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-01-16T07:56:17+00:00","versionOfRecord":[],"versionCreatedAt":"2026-01-13 04:37:40","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8456967","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8456967","identity":"rs-8456967","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}