https://orcid.org/0000-0002-5858-8463
Figures
Abstract
Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae) is a key natural predator of agricultural and forestry pests. In nature, temperature affects the growth, development and predation ability of predators. Therefore, this study assessed the growth, development, and reproduction of E. furcellata at 20, 23, 26, 29, and 32°C. Age-stage, two-sex life table analysis showed that the development duration of each stage decreased with increasing temperature. At 20°C, individuals reached adulthood but females did not oviposit. At 29°C, intrinsic and finite rates of increase and fecundity were higher, with values of 0.12, 1.13 and 41.59, respectively. Moreover, the mean generation time was relatively short at 29.98 d. Spodoptera frugiperda (Smith) (Lepidoptera: Noctuidae) is a highly destructive invasive pest that causes severe economic losses to crops. Therefore, this study evaluated the potential of E. furcellata to control S. frugiperda by predation functional response and interference effects. The functional response of adults to fourth-instar larvae of S. frugiperda followed the Holling II equation across all tested temperatures. Predation ability (a/Th) and maximum daily predation (1/Th) were the highest at 32°C (female:a/Th = 52.7149, 1/Th = 51.8135; male:a/Th = 46.2538, 1/Th = 44.8430), but adult search efficiency was negatively correlated with prey density. At constant prey density, search efficiency increased with temperature. Intraspecific competition and mutual interference were also observed among adults. Across temperatures and prey ratios, adults consistently exhibited strong predation preference for fourth-instar larvae of S. frugiperda. These results provide a theoretical basis for the practical use of E. furcellata in pest management.
Citation: Liu L, Yao M, Zhao R, Chen W (2026) Analysis of temperature adaptability of Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae) based on age-stage, two-sex life table and predatory functional response. PLoS One 21(3): e0344773. https://doi.org/10.1371/journal.pone.0344773
Editor: J Joe Hull, USDA Agricultural Research Service, UNITED STATES OF AMERICA
Received: October 17, 2025; Accepted: February 24, 2026; Published: March 12, 2026
Copyright: © 2026 Liu et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Data Availability: All relevant data are within the manuscript and its Supporting Information files.
Funding: The author(s) received no specific funding for this work.
Competing interests: The authors declare no conflict of interest.
1. Introduction
Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) is a highly destructive pest characterized by broad feeding habits, high fecundity, strong adaptability, and extensive dispersal. The Food and Agriculture Organization of the United Nations categorizes it as a “globally significant agricultural pest with transboundary migratory flights” [1]. Eocanthecona furcellata (Wolff) (Hemiptera: Pentatomidae) is an important predatory natural enemy of agricultural and forestry pests. It preys on more than 40 pest species in the orders Lepidoptera, Coleoptera, Hymenoptera, and Hemiptera [2,3], and is now widely used to control S. frugiperda, Spodoptera. litura (Fabricius) (Lepidoptera: Noctuidae), Plutella xylostella L. (Lepidoptera: Plutellidae), and Helicoverpa assulta (Guenee) (Lepidoptera: Noctuidae), among other species [1,4]. The use of E. furcellata for pest control is efficient and environmentally friendly. At present, the mass rearing of E. furcellata incorporates provenance purification, hierarchical breeding and environmental regulation. However, owing to high breeding cost and environmental sensitivity, challenges in mass rearing and field release persist [5,6].
In nature, temperature is a major abiotic factor influencing the growth, development, and predation ability of natural enemy insects. Moreover, it strongly affects their mass propagation and field use [7–9]. Numerous studies have shown that extreme temperatures markedly affect insect growth and development [10–12]. For example, at 18°C–30°C, increasing temperature shortened the developmental duration, adult longevity, and generation cycle of Arma chinensis (Fallou) (Hemiptera: Pentatomidae) [13]. However, a critical knowledge gap remains regarding how temperature affects the biological traits and predatory performance of E. furcellata. Thus, investigating the growth, development, and predation of E. furcellata across temperatures to assess its developmental duration and control potential can enhance the efficiency of large-scale production during pest outbreaks and improve feasibility and field release efficiency, reduce the use of chemical pesticides, and promote green development. [14,15].
Constructing life tables is a standard and effective method for evaluating insect survival and adaptability at different temperatures [16,17]. Chi and Liu first proposed the age-stage, two-sex life table by accounting for differences between individuals and sexes [18]. Later improvements produced the user-friendly TWOSEX-MSChart, which enables accurate analysis of insect population dynami [19–21]. For example, Xu et al. used temperature as a key factor in evaluating growth and predation in Chrysoperla sinica (Tjeder) (Neuroptera: Chrysopidae) with the age-stage, two-sex life table, showing that spring was the best time to release the species to control Sitobion avenae (Fabricius) (Hemiptera:Aphididae) [9]. Similarly, Dargazani and Sahragard comprehensively evaluated the control potential of Aphis gossypii (Glover) (Hemiptera:Aphididae) by the introduced Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Iran using the age-stage, two-sex life table, showing that H. axyridis strongly suppresses A. gossypii [16].
When evaluating the effect of temperature on natural enemies, considering its influence on predation is essential to maximize biological control. For example, Omkar and Kumar reported that the predation rate of Coccinella transversalis (Fabricius) (Coleoptera: Coccinellidae) and Coccinella septempunctata L. (Coleoptera: Coccinellidae) increased and then declined with increasing temperature from 15°C to 35°C [22]. Fuethermore, Li et al. demonstrated that temperature modulates the instantaneous attack rate and search efficiency of natural enemies. Notably, the instantaneous attack rate of female Scolothrips takahashii (Thysanoptera: Thripidae) increased linearly with rising temperature [23].
The present study assessed E. furcellata at various temperatures: 20, 23, 26, 29, and 32°C. The age-stage, two-sex life table was used to evaluate temperature effects on the species’ growth and development. The effect of temperature on predation of S. frugiperda was also analyzed by fitting predation function and interference responses. Finally, the predation preference experiments were performed to explore whether the feeding of Tenebrio molitor L. (Coleoptera: Tenebrionidae) pupae during the feeding stage affected its predation on S. frugiperda. The results provide a scientific basis for indoor propagation and field release of E. furcellata.
2. Materials and methods
2.1 Insect rearing
Populations of E. furcellata were collected from maize fields at the teaching experimental field of Guizhou University, Guiyang, China (26.41°N, 106.68°E), and were subsequently reared for >10 generations in a climate chamber (58.7 cm × 51.3 cm × 185 cm, Ningbo Jiangnan Instrument Factory, Ningbo, China) with T. molitor pupae at 26°C ± 1°C, 70% ± 5% relative humidity (RH), and a 16:8-h light:dark photoperiod. To maintain genetic diversity, field-collected populations were regularly introduced for genetic renewal. T. molitor pupae were obtained from Weihai Jiulian Biotechnology Co., Ltd. (Shandong, China) and reared on a wheat bran diet. Larvae were maintained at 26°C and 55%–60% RH, fed wheat bran, and reared for 5–6 weeks before pupae were separated from bran and frass. S. frugiperda larvae was collected in June 2023 from a maize field in Longping, Luodian County, Guizhou Province, China (25.48°N, 106.63°E).
2.2 Test methods
Experiments were conducted at five constant temperatures (20, 23, 26, 29, or 32°C) and 70% ± 5% RH under a 16:8-h light:dark photoperiod. At each temperature, E. furcellata was reared in a climate chamber from eggs to adults for subsequent tests.
2.2.1 Effects of temperature on growth, development, and reproduction in E. furcellata.
Groups of 100 eggs (successfully tracked) were placed in climate chambers at each temperature. After the group of eggs hatched, first-instar nymphs were individually placed in feeding boxes (6.5 cm in diameter and 3.5 cm in height) and reared at each temperature. Each instar was marked with a serial number and provided with a wet cotton ball and T. molitor pupae (first instars were given only wet cotton balls, as they do not prey). Sex was noted, and molting, mortality, and eclosion were observed and recorded for each E. furcellata individual daily, with food replenished as needed and feeding boxes cleaned and replaced. After adult emergence, one newly emerged female and one male were paired in a clean feeding box (15.2-cm length, 9.9-cm width, and 6.0-cm height; consistent in the experiments described below) and supplied daily with wet cotton balls and fresh T. molitor pupae. Oviposition date, egg count, and time of death were recorded.
2.2.2 Predatory function and search efficiency of E. furcellata on S. frugiperda at different temperatures.
Based on pre-experiments, density gradients of fourth-instar S. frugiperda larvae were set at 5, 10, 15, and 20 larvae per box. A wet cotton ball and fresh maize leaves were added to each box to reduce S. frugiperda larval cannibalism. One 24 h-starved E. furcellata reared at the corresponding temperature was introduced into each box, which was then placed in a climate chamber at that temperature. Male and female adults were tested separately. Each prey density treatment was repeated five times. After 24 h, the number of surviving prey larvae in each box was recorded.
2.2.3 Effect of E. furcellata density on predation of S. frugiperda at different temperatures.
Based on pre-experiments, E. furcellata adult densities were set at 1, 2, 3, 4, and 5 individuals per box, and each individual was starved for 24 hours before the experiment. Both sexes reared at the target temperatures were used, with males and females tested separately. Each box received 60 fourth-instar S. frugiperda larvae, a wet cotton ball, and sufficient maize leaves to reduce larval self-injury. Boxes were then placed in climate chambers at the corresponding temperatures. Each treatment density was repeated five times. After 24 h, the number of surviving larvae in each box was recorded.
2.2.4 Predation preference of E. furcellata between S. frugiperda and T. molitor at different temperatures.
Based on pre-experiments, three prey ratios were tested: S. frugiperda:T. molitor at 10:20, 15:15, and 20:10 individuals per box. Male and female adults were tested separately. Different ratios of fourth-instar S. frugiperda larvae and T. molitor pupae were placed in feeding boxes, and one 24 h-starved E. furcellata adult reared at each temperature was introduced. Prey were counted after 24 h. Each treatment was repeated five times. All experiments were conducted in artificial climate chambers.
2.3 Data analysis
Parameters of the age-stage, two-sex life table were estimated using the bootstrap method in TWOSEX-MSChart (Chi and Liu 1985) with 100,000 resampled data values [17]. Development time, survival rate, and female daily fecundity were analyzed using this software [24,25]. The same statistical methods were applied to test differences in the age-stage, two-sex life table-related parameters. GraphPad Prism 10 was used to produce histograms and line charts, SigmaPlot 15 was used to create sxj, lx, mx, lxmx, exj, and vxj curves (defined below), and Excel 2021 was used to record and process raw predation-related data. A chi-square test was applied to further validate the predation of E. furcellata against S. frugiperda. Degrees of freedom (df) were calculated as (where Ng represents the number of groups and Np is the number of model parameters), and significance was defined as P < 0.05. When χ² < χ²0.05, df and P > 0.05, the model was considered well fitted, with no significant difference between theoretical values and observed values. The functional response was fitted using the Holling II disk equation. Predation parameters were estimated via the least-squares method, 95% confidence intervals were calculated, and predation charts were created using GraphPad Prism 10.
2.3.1 Formulae for life table parameters.
- (1) The age-specific survival rate (lx), i.e., survival from egg to age x, where Sxj is the age-stage-specific survival rate (probability of E. furcellata surviving from egg to age x stage j), was calculated as follows
- (2) The age-specific fecundity rate (mx), i.e., average fecundity of the E. furcellata population at age x, where fxj is age-specific fecundity in female adults (eggs laid at age x stage j), was determined as follows:
- (3) The population age-specific reproduction value (lxmx), i.e., product of lx and mx, was calculated as follows:
- (4) The age-stage life expectancy (exj), i.e., expected days of survival for E. furcellata at age x stage y, where n is the last population age for each treatment, and s’iy is the probability that an individual of age x stage y survives to age i stage j, was determined using the following formula:
- (5) The age-stage reproductive capacity (vxj), i.e., contribution of an E. furcellata individual at age x stage y to the population’s future, was calculated as follows:
- (6) The intrinsic rate of increase (r), i.e., the maximum instantaneous population growth rate under stable conditions (specific physical and biological conditions), was determined as follows:
- (7) The finite rate of increase (λ), i.e., average daily growth rate of the E. furcellata population without external environmental limitations, was calculated as follows:
- (8) The net reproductive rate (R0), i.e., total offspring produced per individual in a lifetime, was determined using the following formula:
- (9) The average generation period (T), i.e., time required for successive generations under stable growth, was calculated as follows:
- (10) The gross reproduction rate, i.e., the sum of mx values, was determined as follows:
2.3.2 Formula for predation experiment.
The predator–prey functional response was modeled using the Holling II type disk equation:
where Na represents the number of fourth-instar S. frugiperda larvae preyed upon, a is the instantaneous attack rate of E. furcellata toward S. frugiperda, N is the set prey density, Tr is the total duration, and Th represents the handling time of E. furcellata per to S. frugiperda larva [26,27].
The search effect formula is as follows:
where S is the search effect value, and other parameters are defined above [28].
The interference effect formula is as follows:
The Hassell–Verley model was used to fit self-density interference. In the above formula, E is the predation rate, Na the number of prey consumed, N the initial prey count, P the initial predator density, Q the search coefficient, and m the interference coefficient [29].
The selectivity index (D) of E. furcellata toward prey was determined as follows:
where N1 and N2 are initial S. frugiperda and T. molitor densities, respectively, and Np1 and Np2 are the S. frugiperda and T. molitor prey consumed, respectively. When D > 1, predators showed preference for a given prey [30].
3. Results
3.1 Effects of temperature on growth, development, and reproduction in E. furcellata
Between 20°C and 32°C, developmental duration and preadult stages of E. furcellata shortened with increasing temperature. Male and female adults lived significantly longer at 20°C compared with other temperatures. Both the adult preoviposition period and total preoviposition period were shortest at 29°C. Average fecundity increased with temperature, peaking at 32°C but differences from 26°C and 29°C were not significant (Table 1).
At 29°C, R0 reached 41.59, r was 0.12, and λ was 1.13, which are higher than those at other temperatures and significantly higher than at 23°C. T was significantly longer at 20°C compared with the other temperatures. GRR peaked at 26°C but did not differ significantly across treatments (Table 2).
Stage overlap was pronounced in E. furcellata. Except for eggs, survival rates increased and then declined across instars. At 23°C, adults survived longest, with males and females dying at 115 and 106 days, respectively (Fig 1).
Across temperatures, lx declined with E. furcellata age. Additionally, fx and mx fluctuated, showing a pattern of increase, decline, and eventual decrease to 0 (Fig 2). Except for eggs and first-instar nymphs at all temperatures and second to fourth instars at 26°C, exj showed a general trend of decreasing, increasing, and then declining with development time (Fig 3). Similarly, vxj in E. furcellata increased with developmental duration, with peak values increasing at higher temperatures (Fig 4).
3.2 Predatory function and search efficiency of E. furcellata on S. frugiperda at different temperatures
Predation by both E. furcellata sexes on S. frugiperda generally increased with prey density (Fig 5). Chi-square tests (x2 = 1.5, df = 17) are less than critical values (x20.05, 17 = 27.59), then P > 0.05, suggesting no significant difference between theoretical and actual values. Affirming the functional response to fourth-instar S. frugiperda larvae fitted the Holling II model well (Table 3), indicating that E. furcellata predation was significantly correlated with prey density across temperatures. For female E. furcellata, the highest instantaneous attack rate on S. frugiperda (a = 1.0223) occurred at 29°C. Predation ability (a/Th) and maximum daily consumption (1/Th) increased with temperature, peaking at 32°C (52.7149 and 51.8135, respectively). Male predation parameters followed the same trend, with maxima at 32°C (a = 1.0315, a/Th = 46.2538, and 1/Th = 44.8430). Handling time (Th) decreased as temperature increased. Thus, temperature had a significant effect on E. furcellata predation in S. frugiperda: predation efficiency was low at 20°C–23°C and high at 26°C–32°C.
Search efficiency in E. furcellata was negatively correlated with S. frugiperda prey density at all temperatures. At a constant temperature, E. furcellata searching decreased with increasing prey density; at a constant prey density, search efficiency increased with increasing temperature. These results indicate that temperature had a significant effect on the search efficiency of E. furcellata preying on S. frugiperda (Fig 6).
3.3 Effect of E. furcellata density on predation of S. frugiperda at different temperatures
In a defined space, increasing predator density often induces considerable intraspecific interactions, a phenomenon termed mutual interference. This interference reduces the foraging efficiency of individual predators as density increases. The Hassell–Verley interference model was applied to fit the predation rate of E. furcellata adults on fourth-instar S. frugiperda larvae under varying temperatures and predator densities. The correlation coefficient between predation rate and predator density was R2 > 0.97, and chi-square tests values were below the 0.05 critical threshold (χ² < χ²0.05, 17), indicating P > 0.05 and strong agreement between observed and theoretical values. The model effectively characterized the mutual interference of E. furcellata during predation.At constant predator density, E. furcellata predation on fourth-instar S. frugiperda larvae increased with temperature. Higher predator densities increased the total prey killed and scramble competition intensity but reduced the predation rate and per-capita predation, indicating competition and mutual interference among individuals with a specific spatial range. (Tables 4 and 5).
3.4 Predation preference of E. furcellata between S. frugiperda and T. molitor at different temperatures
The predation preference of E. furcellata for fourth-instar S. frugiperda larvae and T. molitor pupae at different temperatures is presented in Tables 6 and 7. When S. frugiperda and T. molitor coexisted in varying proportions, the selectivity index of male and female E. furcellata adults for fourth-instar S. frugiperda larvae was > 1 in all three combinations. This indicates that both E. furcellata sexes exhibited a strong predation preference for fourth-instar S. frugiperda larvae.
4. Discussion
Studying the effects of temperature on the growth, development, and reproduction of natural enemy insects is critical for ecological regulation of field pests and indoor artificial mass-rearing, particularly for improving biological control efficiency [31]. In the present study, at 20°C, eggs hatched and developed into adults but female adults failed to oviposit, consistent with the findings of Zhu [32]. This may be due to prolonged preoviposition periods and shortened female lifespan at 20°C, causing death before egg laying. In addition, Xu et al. reported that egg and nymph development in A. chinensis was significantly shortened as temperature increased from 18°C to 30°C [33]. Similarly, the development of each instar of Picromerus bidens L. and Picromerus lewisi (Fallou) decreased as temperature increased from 18°C to 32°C, and P. bidens eggs failed to hatch normally at 15°C and 35°C [34,35].
Population parameters are key for evaluating treatment effects on insects [36]. The current study found that E. furcellata completed development at 23°C with long survival but low predation, indicating that eggs can be stored at this temperature during winter conservation. Prolonging E. furcellata development at 23°C extends conservation periods and reduces feeding costs. Thus, relatively low temperatures in practical applications can prolong E. furcellata shelf life and reduce mortality due to untimely feeding during transport. At 26°C, the developmental duration of each instar was moderate, population fecundity was high, and adult longevity was prolonged, with E. furcellata completing 8–9 generations annually. Compared with 26°C, r, λ, and fecundity were higher at 29°C, with a shorter generation cycle, indicating that this temperature is more favorable for large-scale propagation of E. furcellata.
For E. furcellata, predation under five constant temperatures followed the Holling II model, consistent with previous studies on natural enemy insects on young S. frugiperda larvae [37–45]. Predation of E. furcellata adults toward S. frugiperda larvae increased with temperature. When the temperature was 32°C, the a/Th and 1/Th of female adults were the highest, which were 52.7149 and 51.8135, respectively. The a/Th and day 1/Th of male adults were also the highest, 46.2538 and 44.8430, respectively,
consistent with studies assessing 17°C–37°C treatments [46]. These adults showed the strongest predation at high temperatures, whereas low temperatures limited activity and control efficiency.
The search efficiency of E. furcellata adults negatively correlated with S. frugiperda prey density at each temperature. Additionally, the prey search efficiency of E. furcellata adults increased with temperature, being lowest at 20°C. At temperatures above 29°C and high prey densities, search efficiency almost plateaued or began to decline, likely due to thermal inhibition of E. furcellata predatory activity, similar to Neoseiulus californicus (McGregor) preying on Eutetranychus chusorientalis (Klein), wherein efficiency decreased above 33°C [47].
Intraspecific competition and mutual interference occurred among adult E. furcellata at the same prey density, increasing with predator density. Search as well as mutual interference coefficients increased with temperature, demonstrating that predation is closely linked to predator density, consistent with the findings of Fan et al. and Tang et al. [1,39]. In the current study, when both fourth-instar S. frugiperda larvae and T. molitor pupae were present, adult E. furcellata consistently preferred S. frugiperda, indicating that feeding T. molitor in the laboratory will not reduce its predation and control of S. frugiperda.
5. Conclusions
In summary, the storage of E. furcellata can effectively prolong its shelf life while reducing rearing costs. Future storage experiments for this species could thus be conducted at approximately 23°C, whereas 29°C favors large-scale propagation. In this study, S. frugiperda was selected as prey owing to its wide distribution, adaptability, rapid reproduction, and severe effects on crops and forests [4]. The current findings support biological pest control concepts, showing that E. furcellata adults effectively prey on fourth-instar S. frugiperda larvae, with the control being strongest at 32°C and weakest at 20°C. Therefore, we speculate that low-temperature releases of E. furcellata require caution, whereas high-temperature summer outbreaks of S. frugiperda can be managed using E. furcellata in the field, the specific results need to be verified by subsequent field experiments. Optimal control should consider actual prey density and appropriate predator release rates. Predation preference experiments confirmed strong adult E. furcellata preference for S. frugiperda, providing guidance for mass propagation and field biological control. Because this study was conducted under constant indoor temperatures, future investigations should incorporate field experiments.
Acknowledgments
The authors express gratitude to Chengxu Wu from the Forestry College of Guizhou University for his support in providing revising the manuscript.
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