Consumption dynamics of Thelosia camina (Schaus, 1896) (Lepidoptera: Apatelodidae) in yerba mateYerba mate (Ilex paraguariensis A. St.-Hil.) is a tree species of the family Aquifoliaceae native to South America with economic, social and environmental relevance in southern Brazil, as it plays a strategic role in the income and maintenance of family farming and in the conservation of remaining Araucaria Forest systems (Antoniazzi et al., 2018; Trevisan, 2023; Souza et al., 2024).
Native and commercial yerba mate are susceptible to several biotic factors, especially defoliating pest insects. Thelosia camina (Schaus, 1896) (Lepidoptera: Apatelodidae), known as yerba mate caterpillar, is considered one of the main pests of the crop, and is often associated with outbreaks of severe defoliation between September and March (Martins et al., 2017; Vidal et al., 2026). Adults have a nocturnal habit and a high oviposition capacity, favoring rapid population replacement under favorable environmental conditions. In addition, the prolonged pupal period in the soil, which can extend for several months, allows the occurrence of seasonal population peaks (Iede and Soares, 2000; Penteado Júnior and Goulart, 2019; Thomazini, 2010).
The management of yerba mate caterpillar presents additional challenges due to the destination of the crop leaves for human consumption, which restricts the application of conventional insecticides and increases the demand for safe and environmentally sustainable control strategies (Górka et al., 2025). Currently, management is based on cultural and mechanical methods, including manual removal of postures, visual monitoring, and capture of adults with light traps.
The temporal organization of herbivory remains poorly understood for most defoliating species of forest importance. Although the daily feeding pattern have been described for Lepidoptera of agricultural importance, there is no information for yerba mate caterpillar. The characterization of these patterns is essential to understand insect-plant interactions, estimate the potential for damage and improve Integrated Management programs. Therefore, this research aimed to characterize the daily feeding pattern and the temporal dynamics of leaf consumption of the yerba mate caterpillar, providing unprecedented information on feeding behavior.
Thelosia camina larvae were collected from a 25-year-old commercial yerba mate plantation in Santa Maria do Oeste, Paraná, Brazil (24°56’ S; 51°51’ W; 1,049 m), on March 8, 2023 and evaluated at the Entomology Laboratory of the State University of the Midwest (UNICENTRO), CEDETEG Campus, Guarapuava, Paraná. The region, inserted in the Third Paraná Plateau, has a humid temperate Cfb (Köppen) climate, with no defined dry season.
Insects from natural populations were collected from more than 100 yerba mate trees, with an average of approximately 20 larvae per tree. Natural populations wereused because the long development cycle, the prolonged pupal period and seasonality limit the maintenance of laboratory colonies (Iede and Soares, 2000; Penteado Júnior and Goulart, 2019; Thomazini, 2010), giving greater ecological relevance to the results. The larvae were acclimatized for 48 h in plastic containers (29 × 19 × 10 cm) with fresh yerba mate leaves, under 25 ± 2 °C, RH of 70 ± 10% and photophase of 14 h The food was then removed, and the larvae were subjected to a 10-h fasting period before the bioassay. Then, fourth - instar larvae were individualized into 20 experimental units (n = 20) in 300 mL containers lined with filter paper and moistened cotton, containing a mature yerba mate leaf (80-120 cm2; mean 100 cm2), previously selected for experimental standardization, the same leaf was used throughout the 24 h evaluation period. (Sanané et al., 2021; Wang et al., 2024).
The bioassay, in a completely randomized design, began at 12:00 h. Leaf intake was quantified every four hours for 24 h (16:00, 20:00, 00:00, 04:00, 08:00 and 12:00 h), under photoperiod of 14L:10D (scotophase at 20:00 h; photophase at 06:00 h). In scotophase, images were obtained under red LED (660 nm; 3-5 W) to minimize interference in the daily feeding pattern. The area consumed was estimated in ImageJ (v. 1.46r) by the difference between consecutive measurements (Ferreira and Rasband, 2012; Schneider et al., 2012).
The data was analyzed in R v. 4.6.0 (R Core Team, 2021). The assumptions of normality and homogeneity of variances were verified. For repeated measures, Mixed Linear Models (LMM) were used, with time (six levels) as a fixed effect and the caterpillar identity as a random intercept effect (Pinheiro and Bates, 2000). The models were fitted with lme4 (Bates et al., 2015) and evaluated by DHARMa residue simulation (Hartig, 2022). The significance of the fixed effects was determined by Type III ANOVA with Kenward-Roger approximation for estimating the degrees of freedom, implemented in the lmerTest package (Kuznetsova et al., 2017) and, when detected (α = 0.05), the estimated marginal means (EMMs) were compared by Tukey's test, using the emmeans package (Lenth, 2024). Leaf consumption rate was estimated using simple linear regression of the mean leaf area consumed over time, reflecting the temporal feeding pattern of the population.
The species T. camina consumed, on average, approximately 15 cm2 of leaf area per caterpillar in 24 hours, indicating high herbivory potential (Figures 11b). Although individual consumption seems limited, its relevance becomes significant considering population outbreaks and the simultaneous feeding of numerous individuals. Continuous removal of photosynthetic tissue affects carbon assimilation, vegetative growth, and post-defoliation recovery (Wang et al., 2021; Zhang et al., 2022). In perennial crops such as yerba mate, successive losses of leaf biomass can compromise leaf replacement and reduce productivity, especially in periods of high population density (Pacheski et al., 2023; Wetzel et al., 2023). However, the magnitude of the effects depends on the frequency and temporal distribution of feeding events, making food chronobiology relevant to understand insect-plant interactions and improve Integrated Pest Management strategies (Galli et al., 2024; Delamore et al., 2026).
Accumulated leaf intake (cm2) (a, b) and leaf area consumed (cm2) at successive intervals of four hours (c) of yerba mate (Ilex paraguariensis) by Thelosia camina larvae for 24 hours, under controlled laboratory conditions (25 ± 2 °C; photoperiod of 14 h). Means followed by the same letter do not differ from each other, according to Tukey’s test applied to estimated marginal means (MMEs), obtained by Mixed Linear Model (LMM) (p ≤ 0.05).
Feeding activity showed a defined temporal pattern (Figure 1c), with higher consumption between 12:00 and 16:00 h and a significant reduction between 00:00 and 04:00 h, demonstrating temporal organization probably associated with the interaction between endogenous circadian mechanisms and environmental factors (Patke et al., 2020; Saunders, 2020). Unlike Spodoptera spp. and Trichoplusia ni, whose consumption is more intense in scotophase, a behavior associated with reduced exposure to visually oriented predators (Bloch et al., 2013; Montezano et al., 2019), T. camina showed higher feeding activity during the daytime under laboratory conditions, indicating a distinct temporal feeding pattern that may be associated with the temporal dynamics of the host plant and environmental conditions (Joo et al., 2019; Erb and Reymond, 2019; Smith et al., 2026). However, the accumulated consumption increased continuously, indicating that momentary oscillations do not determine the final impact of herbivory. In perennial plants, the persistence of feeding throughout development may be more determinant than instantaneous consumption, unlike fast-growing larval species, such as Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera: Noctuidae) Helicoverpa armigera (Hübner, 1808) (Lepidoptera: Noctuidae) and Bedellia somnulentella (Zeller, 1847) (Lepidoptera: Bedelliidae) (Montezano et al., 2019; Bakry and Abdel-Baky, 2023; Volp et al., 2024; Freitas et al., 2026).
Appliedly, the identification of periods of greater activity helps in the planning of sampling and in the timing of application of ingestion-dependent microbiological agents, such as Bacillus thuringiensis Berliner, 1915 (Bacillales: Bacillaceae) and baculovirus (Pardo-López et al., 2013), inserting itself in the perspective of chronoculture (Smith et al., 2026). The diurnal pattern may reflect adaptation to the species' climatic niche, which is abundant between September and March in southern Brazil (Penteado Júnior and Goulart, 2019; Vidal et al., 2026), although tritrophic interactions in the field can modulate this behavior (van Doan et al., 2021; Stell et al., 2022). This study provides the first characterization of the daily leaf consumption of T. camina, demonstrating an organized feeding rhythm with diurnal predominance, high defoliation potential and implications for the targeted temporal management of this pest.
Thelosia camina showed temporally structured feeding activity in yerba mate leaves, with higher consumption between 12:00 and 16:00 h and reduction during the night. Although the data were obtained under laboratory conditions, they still offer subsidies for the formulation of hypotheses applied to monitoring and guided temporal management. Field studies are necessary to verify if the observed pattern reproduces under cultivation conditions.
Acknowledgements
To the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES Finance code - 001).
Data Availability Statement
The research data is available upon prior request via email to the corresponding author.
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e-mail: cmaciel@unicentro.brEditor:
