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Article

Evaluation of Predation on Phytophagous Insects by a Phytozoophagous Mirid Bug, Apolygus lucorum

1
Yantai Academy of Agricultural Sciences, Yantai 265500, China
2
State Key Laboratory for Biology of Plant Diseases and Insect Pests, Institute of Plant Protection, Chinese Academy of Agricultural Sciences, Beijing 100193, China
3
National Center of Technology Innovation for Comprehensive Utilization of Saline-Alkali Land, Dongying 257300, China
*
Author to whom correspondence should be addressed.
Insects 2026, 17(4), 397; https://doi.org/10.3390/insects17040397
Submission received: 2 March 2026 / Revised: 3 April 2026 / Accepted: 4 April 2026 / Published: 7 April 2026
(This article belongs to the Special Issue Biosystematics and Management of True Bugs (Hemipterans))

Simple Summary

Apolygus lucorum is a significant agricultural pest that also exhibits predatory behavior on small arthropods. This study systematically evaluated its predatory capacity against eggs of Helicoverpa armigera, nymphs of Aphis gossypii, and nymphs of Bemisia tabaci. Laboratory experiments demonstrated that A. lucorum showed the strongest predatory preference for A. gossypii. Its predatory ability increased with developmental stage, and males generally exhibited higher predation efficiency than females. DNA-based gut content analysis of A. lucorum in Bt cotton fields confirmed its predation on A. gossypii in natural environments. These findings systematically elucidate the predatory characteristics and ecological adaptability of A. lucorum, contributing to a deeper understanding of its functional role in agricultural ecosystems.

Abstract

Apolygus lucorum, a phytozoophagous mirid bug, plays an important role in the species interactions within fruit tree and cotton ecosystems. Previous research has mainly focused on the phytophagous damage that it causes to crops, while its role as a predator of arthropods remains poorly understood. In this study, we systematically investigated the functional responses of A. lucorum to three crop pests: eggs of Helicoverpa armigera, nymphs of Aphis gossypii, and nymphs of Bemisia tabaci. The results show that the predatory behavior of A. lucorum towards all three prey species followed a Holling type II functional response model. Predatory performance varied significantly depending on prey species, developmental stage, and sex of the mirid. The theoretical maximum predation rate was highest for A. gossypii (833.33 individuals/day) and lowest for B. tabaci nymphs. Adult mirids and older nymphs (4th instar) exhibited higher predation rates than younger nymphs. Field-collected A. lucorum from Bt cotton fields were analyzed using molecular diagnostics, and the result confirmed natural predation on A. gossypii, which was consistent with observed pest occurrence patterns in the field. Overall, this study clarifies the prey selectivity and stage-dependent predatory strategies of A. lucorum, providing insights into its trophic flexibility as a facultative predator. These findings contribute to a more comprehensive understanding of its ecological role in agricultural ecosystems, but do not support its use as a biological control agent given its predominantly phytophagous nature and documented pest status.

Graphical Abstract

1. Introduction

Apolygus lucorum (Hemiptera: Miridae) is a phytozoophagous pest widely distributed in agricultural ecosystems, with over 200 recorded host plants. It causes significant damage to cotton and several fruit crops including jujube, grape, and peach [1,2,3,4]. Traditionally, A. lucorum is unequivocally defined as a phytophagous pest. Both adults and nymphs of A. lucorum feed by piercing tender plant tissues such as buds, young leaves, flower buds, and immature fruits. This feeding behavior initially results in small dark-brown necrotic spots that progressively expand and coalesce as plant development continues, ultimately leading to leaf perforation, fruit deformity, and bud abscission. These symptoms severely compromise crop yield and quality, frequently resulting in substantial economic losses [5,6,7,8]. Consequently, research has predominantly focused on the population dynamics, damage mechanisms, and control strategies of A. lucorum [9,10,11].
However, emerging evidence from field observations and laboratory studies indicates that A. lucorum exhibits facultative predatory behavior. In addition to feeding on plants, it actively preys on small arthropods such as aphids, whiteflies, and lepidopteran eggs [12,13]. This mixed feeding strategy—primarily phytophagous with opportunistic zoophagy—is not uncommon among mirid bugs. For example, the tarnished plant bug, Lygus lineolaris (Hemiptera: Miridae), a major pest of cotton, alfalfa, and strawberry, has also been reported to prey on spider mites and aphids [14,15]. Similarly, Adelphocoris suturalis (Hemiptera: Miridae), in addition to its phytophagous damage to cotton, can consume eggs and early-instar larvae of Spodoptera exigua (Lepidoptera: Noctuidae) and Agrotis ipsilon (Lepidoptera: Noctuidae) [16]. Feng et al. [17] further documented that adults of Adelphocoris lineolatus (Hemiptera: Miridae) can prey on early-instar larvae of Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Collectively, these observations suggest that facultative predation represents a widespread ecological adaptation within the Miridae family, likely enhancing population stability and ecological flexibility in heterogeneous agricultural environments.
From an ecological perspective, insects possessing facultative predatory traits often function as key species in agroecosystems, contributing to the maintenance of community stability and the regulation of trophic interactions. In primarily phytophagous mirids, the expression of zoophagy is generally considered a nutritionally driven strategy, often triggered by the need for specific nitrogenous compounds or by suboptimal plant resources [18]. For A. lucorum, predation may occur when prey availability aligns with periods of high nutritional demand, such as during nymphal development or adult reproduction, or when preferred host plant tissues become scarce [19,20]. These insects not only directly affect prey populations through predation but may also indirectly influence plant performance and community assembly [21,22,23]. For instance, in protected cultivation systems, mirid species such as Nesidiocoris tenuis (Hemiptera: Miridae) and Macrolophus pygmaeus (Hemiptera: Miridae) have proven effective in suppressing populations of whiteflies, thrips, and spider mites, thereby modulating their seasonal abundance patterns [24,25,26]. Such findings provide important insights into the ecological functions of facultatively predatory mirids.
A. lucorum is one of the most widely distributed and economically damaging omnivorous mirid pests in Chinese agroecosystems. Its population has increased significantly following the large-scale adoption of Bt cotton, and it has become a major pest of cotton, jujube, grape, and other crops [2,3]. In China, A. lucorum populations are characterized by high densities and prolonged seasonal activity, a trend exacerbated by the expansion of horticultural crops such as fruit trees and reduced insecticide applications in Bt cotton fields, collectively driving a substantial rise in its abundance and amplifying its ecological and economic influence within agricultural ecosystems [27,28]. Understanding the conditions that trigger predation in A. lucorum is therefore critical for predicting its behavior in field settings.
Given the documented pest status of A. lucorum and the increasing recognition of its facultative predatory behavior, this study aims to characterize its predation capacity under controlled conditions while critically evaluating the extent to which such capacity translates to natural field settings. By integrating laboratory functional response assays with field molecular detection, we seek to determine whether predation by A. lucorum represents a functionally significant regulatory force or merely an opportunistic nutritional supplement. This distinction is essential for accurately interpreting its ecological role and avoiding overestimation of its potential as a natural enemy in integrated pest management (IPM) programs.
In this study, we investigated the predatory potential of A. lucorum toward three economically important pests in China: eggs of Helicoverpa armigera (Lepidoptera: Noctuidae), nymphs of Aphis gossypii (Hemiptera: Aphididae), and nymphs of Bemisia tabaci (Hemiptera: Aleyrodidae). Under controlled laboratory conditions, we systematically quantified the functional response of A. lucorum to these prey types, with particular emphasis on the effects of predator developmental stage (nymphal instars and adults), sex, prey species, and prey density on key foraging parameters including instantaneous attack rate, handling time, and estimated maximum daily consumption. In parallel, field surveys were conducted in cotton fields to monitor the seasonal dynamics of A. lucorum and its potential prey. Molecular gut content analysis was employed to detect predation events in field-collected A. lucorum individuals, providing direct evidence of its feeding on pests such as A. gossypii under natural conditions. By integrating laboratory experiments with field validation, this study aims to establish a comprehensive understanding of the trophic ecology of A. lucorum, thereby offering a scientific basis for evaluating its functional role in agricultural ecosystems and informing the development of ecologically based integrated pest management strategies.

2. Materials and Methods

2.1. Insect Rearing

The A. lucorum colony was established from adults collected in Langfang, Hebei Province, and maintained continuously in the laboratory on green beans sterilized with 5% sodium hypochlorite [29]. Nymphs were reared in transparent plastic containers (19.5 cm × 13.4 cm × 7.2 cm) with fresh or split green bean pods, at a 200–300 density of individuals per container. Several folded filter paper strips (1 cm × 20 cm) were placed at the bottom of each container to facilitate insect movement. The container opening was covered with medical gauze and secured with the lid to prevent escape, and a square opening (7 cm × 7 cm) was cut in the lid for ventilation. Upon reaching adulthood, adults were also fed with green beans, and the ends of the beans were cut at a slant to serve as an oviposition substrate. A cotton ball soaked with 10% honey–water solution was placed on the gauze cover to provide supplement nutrition. After oviposition, green beans bearing egg masses were placed in a ventilated area for 4–5 days and then transferred to clean containers. Egg hatching was monitored daily. Newly hatched nymphs from the same day were transferred to another clean container with fresh green bean pods and reared as described above for subsequent generations. Rearing conditions were maintained at 26 ± 1 °C, 70 ± 5% (RH), and a photoperiod of 14 L/10 D.
The H. armigera colony was established from individuals collected in cotton fields in Xinxiang, Henan Province, and maintained in the laboratory on an artificial diet. Adults were provided with 10% sugar solution as a nutritional supplement. Rearing methods followed Liang et al. [30]. Rearing conditions were 26 ± 1 °C, 70 ± 5% (RH), and a photoperiod of 16 L/8 D.
The experimental populations of A. gossypii and B. tabaci were collected from cotton fields in Langfang and were used directly in experiments without laboratory rearing.

2.2. Laboratory Predation Assays

To comprehensively evaluate the predatory capacity of A. lucorum across different developmental stages, we selected the 2nd instar nymphs, 4th instar nymphs, and 5-day-old adults (both females and males) for the predation assays. The 2nd instar represents the early instar stage with relatively weak predatory capacity, serving as a baseline for developmental comparison. The 4th instar represents the late instar stage, during which rapid growth and high nutritional demands may enhance predatory activity. The 5-day-old adults correspond to the peak reproductive stage, allowing assessment of predatory capacity during periods of elevated nutritional requirements.
To simulate the natural feeding behavior of A. lucorum, no starvation treatment was applied prior to the predation assays. All experiments were conducted under the same environmental conditions as insect rearing (26 ± 1 °C, 70 ± 5% RH, 14 L/10 D photoperiod). Each treatment was replicated five times.
(a)
Predation on H. armigera eggs: Cotton gauze pieces (3 cm × 3 cm) containing H. armigera eggs (laid within 24 h) were prepared. Non-viable or misshapen eggs were removed under a stereomicroscope using an insect pin, and the number of eggs on each piece was recorded. Each gauze piece was placed in a glass tube (7.5 cm × 2 cm), and one A. lucorum individual (2nd instar nymph, 4th instar nymph, 5-day-old female, or 5-day-old male) was introduced using a soft brush. The tube opening was covered with an 80-mesh nylon net and secured with a rubber band to prevent escape. A moistened cotton ball was placed on the net to provide moisture. Prey densities tested were 5, 10, 20, 30, 40, and 50 eggs per tube, with five replicates per density. After 24 h, the number of consumed eggs was recorded under a stereomicroscope based on their shriveled, flattened appearance. Any eggs that had hatched within 24 h (which did not occur) or not shriveled were excluded from consumption counts.
(b)
Predation on A. gossypii nymphs: Cotton leaves infested with A. gossypii nymphs were collected from the field and brought back to the laboratory. Under a stereomicroscope, healthy, unparasitized, and uninfected aphid nymphs (without distinction of instar to better reflect natural field conditions) were carefully selected using a fine brush and transferred onto cotton gauze pieces (3 cm × 3 cm). Each gauze piece containing the designated number of aphids was placed in a glass tubes (7.5 cm × 2 cm), and one A. lucorum individual (2nd instar nymph, 4th instar nymph, 5-day-old female, or 5-day-old male) was introduced into each tube. The tube opening was covered with an 80-mesh nylon net and secured with a rubber band to prevent escape, while a moistened cotton ball was placed on the net to provide moisture. Prey densities tested were 2, 4, 8, 12, 16, and 20 nymphs per tube, with five replicates per density. After 24 h, the predator was removed, and the number of consumed aphids (identified by their darkened, shriveled appearance) was recorded.
(c)
Predation on B. tabaci nymphs: Cotton leaves infested with B. tabaci nymphs were collected from the field and brought back to the laboratory. The leaves were cut into leaf discs (3 cm × 3 cm). Under a stereomicroscope, excess nymphs, as well as unhealthy, parasitized, or infected individuals, were removed, leaving only healthy 4th instar nymphs to ensure uniformity of prey size. Each leaf disc was placed in a glass tube (7.5 cm × 2 cm), and one A. lucorum individual (2nd instar nymph, 4th instar nymph, 5-day-old female, or 5-day-old male) was introduced into each tube. The tube opening was covered with an 80-mesh nylon net and secured with a rubber band to prevent escape, while a moistened cotton ball was placed on the net to provide moisture. The prey densities tested were 2, 4, 8, 12, 16, and 20 nymphs per tube, with five replicates per density. After 24 h, the number of consumed nymphs (identified by their shriveled appearance) was recorded.

2.3. Field Detection of Predation by A. lucorum

2.3.1. COI-Based Molecular Detection Method

(a)
DNA template preparation: Genomic DNA was extracted from individual A. lucorum using a Blood/Cell/Tissue Genomic DNA Extraction Kit (Tiangen Biotech, Beijing, China) following the manufacturer’s instructions. The DNA extracts were stored at −20 °C until use. For PCR amplification, 1 µL of DNA solution was used as the template.
(b)
Species-specific primers: Species-specific primers were designed based on mitochondrial COI gene sequences from H. armigera (GenBank accession No. AY264944) and A. gossypii (GenBank accession No. AY842502). Primers were synthesized by Sangon Biotech (Shanghai, China) and purified by PAGE.
Primers for H. armigera:
Forward primer MLCF: 5′-GGTGATCCTATTTTATATCAC-3′
Reverse primer MLCR: 5′-GAGTATCAATATCTATACCAG-3′
Amplicon length: 239 bp.
Primers for A. gossypii:
Forward primer MYF: 5′-TTCACATCAGCAACTATAATC-3′
Reverse primer MYR: 5′-ACTACATAATAAGTGTCATGC-3′
Amplicon length: 208 bp.
(c)
PCR amplification: PCR reactions were performed in a thermal cycler (Bioer, Hangzhou, China). Each 25 µL reaction mixture contained 0.25 µL of EX Taq polymerase (TaKaRa, Dalian, China), 2.5 µL of 10× PCR buffer (supplied with the Taq polymerase), 2 µL of dNTPs (Tiangen Biotech, Beijing, China), 1 µL of DNA template, 0.5 µL of each forward and reverse primer, and sterile distilled water to a final volume of 25 µL. Each sample was run in duplicate to ensure reliability.
The PCR cycling program was as follows: initial denaturation at 94 °C for 5 min; 36 cycles of 94 °C for 30 s, 55 °C for 30 s, and 72 °C for 1 min; final extension at 72 °C for 10 min.
Positive controls (DNA from H. armigera eggs or A. gossypii nymphs) and negative controls (sterile water instead of DNA template, and DNA from A. lucorum that had not fed on H. armigera eggs or A. gossypii) were included.
(d)
Amplicon detection: A 10 µL of each PCR product was electrophoresed on a 1.2% agarose gel containing 0.5 µg/mL GoldView nucleic acid stain (Solarbio, Beijing, China) at 180 V for 15 min. A DL2000 DNA molecular weight marker (TaKaRa, Dalian, China) was used as a reference. After electrophoresis, the gel was visualized and photographed under UV light using a gel imaging system (Tanon, Shanghai, China).

2.3.2. Field Investigation and Sampling

Field surveys were conducted in cotton fields in Xinxiang City, Henan Province, Xiajin County, Shandong Province, and Langfang City, Hebei Province during the occurrence period of A. lucorum (from late June to early September) in 2009 and 2010. A diagonal five-point sampling method was used, with 10 cotton plants randomly examined at each sampling point, totaling 50 plants per survey site. Surveys were carried out every 3–5 days, and the number of A. lucorum adults, H. armigera eggs, and A. gossypii nymphs per 50 plants were recorded.
In addition, adult A. lucorum were collected during three key periods: the early occurrence stage (late June to early July), the peak occurrence stage (mid-July to early August), and the late occurrence stage (mid-August to early September). Adults were collected using insect nets or aspirators, with as many individuals as possible collected. Collected specimens were placed separately into 2 mL centrifuge tubes containing 100% ethanol, transported to the laboratory within 1 h, then stored at −20 °C or −80 °C for subsequent DNA extraction and PCR analysis.

2.4. Statistical Analysis

All data were analyzed using SAS 9.4 (SAS Institute Inc., Cary, NC, USA), and functional response curve fitting was conducted using R 4.1.3 (R Core Team, Vienna, Austria). A significance level of p < 0.05 was used for all statistical tests. All data are presented as mean ± standard error of the mean (SE).
One-way ANOVA with Tukey’s HSD test (SAS 9.4) was used to compare prey consumption among different developmental stages (2nd instar nymphs, 4th instar nymphs, 5-day-old females, and 5-day-old males) of A. lucorum at each prey density for each prey species (H. armigera eggs, A. gossypii nymphs, and B. tabaci nymphs). Five replicates were conducted per density. Normality and homogeneity of variance were verified using Shapiro–Wilk and Levene’s tests, respectively.
To determine the functional response type, logistic regression was performed on the proportion of prey consumed (Na/N) as a function of initial prey density (N). The model was fitted using a binomial distribution with a logit link function, weighted by initial prey density:
Na/N = exp (P0 + P1 N + P2 N2 + P3 N3)/1 + exp (P0 + P1 N + P2 N2 + P3 N3)
The signs and significance of P1 and P2 were used to determine the functional response type. A Type II response is indicated by a significantly negative linear coefficient (P1 < 0, p < 0.05), whereas a Type III response is indicated by a significantly positive linear coefficient and a significantly negative quadratic coefficient (P2 < 0, p < 0.05). For combinations where P1 was not significant, the functional response type was further evaluated using Holling’s disc equation, with R2 was used as a complementary criterion [31,32].
The predation data conformed to the Holling Type II disc equation [33]:
Na = aTN/(1 + aThN)
where Na is the number of preys consumed, N is the initial prey density, a is the attack rate, T is the experimental time (T = 1 d), and Th is the handling time. Maximum daily prey consumption was calculated as 1/Th, and predation capacity as a/Th. The equation was linearized as
1/Na = 1/aT ×1/N +Th/T
For the population dynamics data, differences in pest densities (A. lucorum adults, H. armigera eggs, and A. gossypii nymphs) among sampling dates were analyzed separately for each location and year using repeated measures ANOVA, followed by Tukey’s HSD test for multiple comparisons. Normality assumptions were tested using the Shapiro–Wilk test.
For the molecular detection data, the positive detection rates of A. gossypii DNA were analyzed separately for each region and year using repeated measures ANOVA. Tukey’s HSD test was used for multiple comparisons. Normality assumptions were tested using the Shapiro–Wilk test. Detection rates of H. armigera eggs were zero across all samples and were therefore excluded from statistical analysis.

3. Results

3.1. Functional Response Type

Logistic regression analysis was employed to determine the functional response types of different developmental stages of A. lucorum to three prey species. The proportion of prey consumed (Na/N) was modeled as a function of initial prey density (N), and the signs and significance of the linear coefficient (P1) and quadratic coefficient (P2) were used to identify the functional response type. Across all predator–prey combinations, the linear coefficient (P1) was consistently negative (Table 1), indicating that the proportion of prey consumed decreased with increasing prey density—a characteristic pattern of a Type II functional response.
Among the 12 predator–prey combinations, only the combination of H. armigera eggs with 5-day-old female adults yielded a statistically significant linear coefficient (P1 = −0.6754, p = 0.0004). For all other combinations, the linear coefficients were negative but not significant (p > 0.05). The quadratic coefficients (P2) were positive across all combinations, which does not satisfy the criteria for a Type III functional response (P1 > 0 and P2 < 0). To further validate the functional response type, nonlinear regression was performed using Holling’s disc equation. The Holling Type II model exhibited excellent goodness-of-fit for all predator–prey combinations, with R2 values ranging from 0.9029 to 0.9965 (Table 1). Taken together, the results from both logistic regression and Holling Type II model fitting confirm that all developmental stages of A. lucorum (2nd instar nymphs, 4th instar nymphs, 5-day-old females, and 5-day-old males) exhibit a Type II functional response when preying on H. armigera eggs, A. gossypii nymphs, and B. tabaci nymphs.

3.2. Predation Rates of A. lucorum on Three Prey Types

The predation rates of A. lucorum at different developmental stages increased with prey density in a decelerating manner and tended to approach saturation at higher densities (Figure 1). The functional response parameters, including attack rate (a), handling time (Th), maximum daily consumption (1/Th), and predation capacity (a/Th), varied considerably among prey types and predator developmental stages (Table 2).
For H. armigera eggs, 2nd instar nymphs reached their maximum consumption (13.00 eggs) at a density of 40 eggs, with no significant difference compared to consumption at 50 eggs (11.20 eggs, p > 0.05). In contrast, the maximum predation rates of 4th instar nymphs, 5-day-old females, and 5-day-old males all occurred at the highest egg density (50 eggs), with values of 31.00, 30.20, and 35.00 eggs, respectively. Across all densities, 2nd instar nymphs consistently exhibited the lowest predation levels (Figure 1A–D). Parameter estimates further supported these observations: 4th instar nymphs and 5-day-old males had the highest predation capacities (a/Th = 75.71 and 73.13, respectively), while 2nd instar nymphs showed the lowest (13.08), consistent with their lower consumption across densities (Table 2).
On A. gossypii nymphs, the predation rate of 2nd instar nymphs fluctuated, peaking at densities of 16 and 20 nymphs (both 6.80 individuals). Consumption at a density of 8 nymphs (5.00 individuals) was higher than at densities of 2, 4, and 12 nymphs. For 4th instar nymphs, 5-day-old females, and 5-day-old males, predation rates increased steadily with aphid density, reaching maxima at 20 nymphs (12.20, 13.20, and 13.60 nymphs, respectively). Across all densities, 5-day-old males consistently consumed more than 5-day-old females. Fourth instar nymphs generally outperformed 2nd instar nymphs, except at the lowest density (2 nymphs) where consumption was lower, and at 8 nymphs where they were comparable (Figure 1E–H). The functional response parameters highlighted these stage-specific differences: 5-day-old males exhibited the highest predation capacity (a/Th = 679.42) and theoretical maximum daily consumption (833.33 aphids), followed by 5-day-old females (273.24 and 400.00 aphids), while 2nd instar nymphs had the lowest values (8.52 and 7.97 aphids) (Table 2).
In assays with B. tabaci nymphs, predation rates generally increased with prey density. Fourth instar nymphs exhibited the strongest functional response, displaying a rapid increase in consumption across the density range—particularly a sharp rise from 12 to 16 nymphs (from 1.90 to 4.00 nymphs)—and reaching the highest maximum consumption (3.60 nymphs) at a density of 20 nymphs. Five-day-old females and males showed more gradual increases without clear inflection points, attaining maxima of 2.70 and 2.90 nymphs, respectively, also at the highest density. In contrast, 2nd instar nymphs reached saturation earlier, with consumption stabilizing at 1.30–1.40 nymphs after a density of 12 nymphs, indicating a lower predation ceiling and earlier saturation. At identical prey densities, 4th instar nymphs consistently outperformed other stages, while 2nd instar nymphs exhibited the lowest predation across the entire density range (Figure 1I–L). Parameter estimates showed that predation capacity for B. tabaci was substantially lower than for the other two prey species. Fourth instar nymphs had the highest a/Th (4.48) and maximum consumption (16.75 nymphs), whereas 2nd instar nymphs again showed the lowest values (0.58 and 3.47 nymphs) (Table 2).
Overall, the functional responses of A. lucorum at different developmental stages to all three prey species conformed to the Holling type II model. Predation capacity and maximum daily consumption increased with developmental stage, with 2nd instar nymphs consistently showing the lowest efficiency and 4th instar nymphs and adults exhibiting significantly higher predatory performance. Notably, 5-day-old males consistently outperformed females across most prey types and densities, suggesting a sex-based difference in foraging strategy, particularly evident in the high predation capacity observed on A. gossypii.

3.3. Detection of Prey Predation by A. lucorum in Cotton Fields

The predatory interactions of A. lucorum with H. armigera eggs and A. gossypii were evaluated using COI molecular markers on individuals sampled from Bt cotton fields. Analysis indicated no positive predation signals for H. armigera eggs in any tested samples from 2009 and 2010. In contrast, predation on A. gossypii by A. lucorum was observed, though with pronounced regional variation. In Langfang, positive detections of A. gossypii predation were recorded consistently across both study years. In 2009, 14 positive samples were identified (5 collected on 15 July and 8 on 18 July), with an average detection rate of 7.04% ± 3.73. In 2010, 8 positive samples were collected between 25 July and 2 August, with an average detection rate of 3.70% ± 2.47. In Xiajin, three positive samples obtained on 27 July 2009, corresponded to an average detection rate of 1.22% ± 1.26, whereas no positive signals were detected in 2010. In Xinxiang, no A. gossypii DNA was detected in either year, resulting in a 0% positive detection rate across both sampling periods (Table 3).
To elucidate the relationship between molecular detection results and actual field ecology, positive detection rates were systematically analyzed alongside the population dynamics of A. lucorum, H. armigera eggs, and A. gossypii during sampling. Positive detections for A. gossypii consistently coincided with peaks in aphid populations, which also corresponded to periods of higher A. lucorum density. In Langfang, positive detections in 2009 corresponded to sampling dates on 15 July, 18 July, and 26 August, with A. gossypii densities of 45,460.00 ± 5480.09, 32,500.00 ± 3904.92, and 1522.00 ± 715.78 individuals per 100 plants, respectively (Figure 2A). In 2010, positive detections were recorded on 25 July, 29 July, and 2 August, accompanied by A. gossypii densities of 12,596.00 ± 1018.56, 23,917.00 ± 2589.97, and 6792.00 ± 657.56 individuals per 100 plants, respectively (Figure 2B). In Xiajin, all three positive reaction samples in 2009 were collected on July 27, when A. gossypii density was 27,760 ± 3757.31 individuals per 100 plants. This date coincided with both the peak occurrence of A. gossypii and the population peak of A. lucorum (Figure 2C). In 2010, despite sufficient sampling effort, the A. gossypii population remained low, with a peak density of only 1246.00 ± 129.38 individuals per 100 plants, and no positive detections were obtained (Figure 2D). In Xinxiang, although the occurrence periods of A. gossypii and A. lucorum broadly overlapped in 2009, the population density of A. lucorum was low and the sample size was limited, resulting in no positive reactions (Figure 2E,F). In 2010, both A. gossypiis and A. lucorum occurred only sporadically, and again no positive reactions were detected (Figure 2F).
As shown in Figure 2, H. armigera egg populations in Bt cotton fields remained at consistently low densities throughout the study period. The highest recorded egg count across all three regions and both years was only 230.00 ± 76.85 eggs per 100 plants. During peaks in H. armigera egg occurrence, A. lucorum was consistently in its early occurrence period in the cotton fields, characterized by low overall population abundance and limited sample availability for molecular analysis. Consequently, no target H. armigera DNA fragments were amplified from any tested A. lucorum individuals.
Analysis based on COI molecular markers indicated that under natural field conditions, A. lucorum did not exhibit significant predation on H. armigera eggs, whereas it showed detectable predation on A. gossypii. Moreover, predation rates varied markedly among geographical regions, with the most substantial predatory activity observed in Langfang. These findings suggest that the feeding behavior of A. lucorum in agroecosystems may be shaped by both spatiotemporal dynamics and prey selectivity.

4. Discussion

This study systematically evaluated the predatory functional responses of A. lucorum at different developmental stages (2nd instar, 4th instar nymphs, and 5-day-old male and female adults) to H. armigera eggs, A. gossypii nymphs, and B. tabaci nymphs. The results indicated that the predatory behavior of A. lucorum towards all three prey types conformed to the Holling type II functional response model, with predation initially increasing and then stabilizing as prey density rose. This pattern aligns with observations of N. tenuis predation of B. tabaci, Calvia muiri (Coleoptera: Coccinellidae) on Panaphis juglandis (Hemiptera: Aphididae), and Arma chinensis (Hemiptera: Pentatomidae) on Spodoptera frugiperda (Lepidoptera: Noctuidae) [34,35,36,37], suggesting that such functional responses are common among predatory insects.
The study further revealed significant influences of developmental stage and sex on predation capacity. When preying on H. armigera eggs, 5-day-old males exhibited the highest predation potential, with a theoretical maximum of 73.53 eggs, followed by 4th instar nymphs (68.97 eggs), while 2nd instar nymphs showed the weakest predation capacity (41.84 eggs). For A. gossypii nymphs, predation differences among stages were more pronounced: 5-day-old males consumed the most (833.33 individuals), significantly higher than 5-day-old females (400.00 individuals), with 2nd instar nymphs consuming the least (7.97 individuals). When preying on B. tabaci nymphs, 4th instar nymphs displayed the strongest predation capacity (16.75 individuals), followed by 5-day-old males (5.74 individuals), while 5-day-old females consumed the least (3.42 individuals). Overall, older nymphs and adults showed a preference for A. gossypii, whereas younger nymphs exhibited a relative preference for H. armigera eggs, indicating that the predatory strategy of A. lucorum is significantly influenced by both its developmental stage and prey type. Similar phenomena have been reported in other predatory insects. For instance, when S. frugiperda and Rhopalosiphum padi (Hemiptera: Aphididae) coexist, Harmonia axyridis (Coleoptera: Coccinellidae) preferentially predates the latter [38]. Further, Cai et al. [39] found that 2nd instar larvae and female adults of H. axyridis exhibited higher predation rates on Aphis spiraecola (Hemiptera: Aphididae) than on Semiaphis heraclei (Hemiptera: Aphididae), while 3rd instar larvae and male adults showed a stronger preference for S. heraclei. These findings collectively highlight the widespread interaction between developmental stage and prey type in predatory insect behavior.
From a developmental perspective, 4th instar nymphs exhibited greater predation capacity than 2nd instar nymphs, indicating enhanced predatory ability with advancing developmental stage. This trend aligns with findings on N. tenuis, whose predation on B. tabaci and Thrips palmi (Thysanoptera: Thripidae) increases with nymphal stage [40]. Regarding sex differences, males consumed more of all prey types than females. This is consistent with observations of Orius albidipennis (Hemiptera: Anthocoridae), where males displayed significantly higher attack rates on Aphis fabae (Hemiptera: Aphididae) than females [41]. This may be related to adaptive behavioral strategies developed by males in mating competition and resource acquisition, with stronger predation capacity likely conferring a competitive advantage within the population [42].
Notably, the predation characteristics of A. lucorum, which are significantly regulated by developmental stage, prey type, and density, are also observed in other mirid species that are primarily phytophagous but facultatively predatory. For example, A. suturalis tends to feed on a mix of plant and animal materials, and its predation on A. ipsilon and S. exigua follows the Holling type II functional response [16,43]. Similarly, Lygus pratensis (Hemiptera: Miridae) stages exhibit enhanced predation on A. gossypii with increased aphid density and nymphal stage, which was further confirmed by field cage experiments [44]. These species demonstrate strong dependence of predatory behavior on prey population density and availability, suggesting that facultative predation may represent an important ecological plasticity for adaptation to agricultural ecosystems.
Although laboratory experiments confirmed the predation capacity of A. lucorum, molecular detection and field population dynamics analyses jointly indicate significant limitations to its actual predatory role under natural conditions. Specifically, A. lucorum consistently failed to exhibit effective predation on H. armigera eggs in the field, while its predation on A. gossypii showed marked regional variation and was strictly concentrated during peak aphid population periods. These contrasting results, on one hand, support the laboratory findings of highest predation capacity on A. gossypii, indicating clear prey preference and population dependence; on the other hand, they reveal that predation on H. armigera eggs is largely unfeasible in the field, as it is constrained by extremely low prey density, high search costs, and phenological asynchrony. These findings underscore that the predatory behavior of A. lucorum is not only influenced by prey type but also closely linked to the spatiotemporal dynamics and population density of prey in the field, highlighting its resource dependence and ecological plasticity as a facultative predator. Consistent with this view, field predation events were detected only during periods of high prey density and were absent when prey populations were low, indicating that the laboratory-derived maximum consumption rates (e.g., 833 aphids/day) are rarely realized under natural conditions due to ecological constraints such as low prey availability, phenological asynchrony, and the pest’s dominant phytophagous behavior.
The transition between phytophagy and zoophagy in A. lucorum appears to be governed by a combination of physiological demands and ecological context. Laboratory results showed that older nymphs and adult males exhibited the highest predation rates, suggesting that periods of rapid growth or reproduction may increase the demand for protein-rich animal prey—a pattern consistent with observations in other omnivorous mirids such as N. tenuis and M. pygmaeus [45,46]. In contrast, under field conditions, such predatory activity becomes evident only when prey are abundant, further supporting the view that zoophagy in this species is predominantly opportunistic rather than obligate. Moreover, its well-documented host-switching behavior—moving among jujube, cotton, and other crops in response to plant phenology—indicates that plant resources remain the primary driver of its population dynamics [47,48]. Thus, while facultative predation may provide nutritional benefits that enhance fitness under specific conditions, it does not alter its fundamental status as a phytophagous pest.
Analyzed in conjunction with its seasonal host-switching behavior, A. lucorum tends to shift towards flowering plants during host transitions [49,50]. When cotton growth enters peak flowering in July, A. lucorum migrates into cotton fields and initiates damage. This migration pattern results in spatiotemporally limited predatory behavior in the field, occurring sporadically only under specific conditions—such as during cotton peak flowering and when high densities of specific prey (e.g., A. gossypii) are present. Although A. lucorum may transiently regulate populations of minor pests like A. gossypii within food webs, its phytophagous damage remains dominant. Predation is merely a supplementary component of its nutritional strategy and does not alter its ecological role as a major pest. Therefore, while the laboratory findings confirm the facultative predatory capacity of A. lucorum, the field evidence demonstrates that this capacity does not translate into reliable biological control under natural conditions. Within an IPM framework, management strategies should continue to prioritize the suppression of A. lucorum as a direct pest, particularly during its peak occurrence periods and host-switching phases. The predatory behavior documented in this study should not be interpreted as a justification for conserving or augmenting A. lucorum populations, as such approaches would risk exacerbating its phytophagous damage. Instead, these findings contribute to a more nuanced ecological understanding of a dominant pest species, informing risk assessment and supporting the development of multi-trophic management strategies that account for complex species interactions within agricultural ecosystems.

5. Conclusions

By integrating laboratory functional response assays, molecular detection, and field ecological surveys, this study systematically clarifies the ecological characteristics and realizes function of the facultative predation behavior exhibited by the primarily phytophagous pest A. lucorum. The results indicate that predation by A. lucorum on H. armigera eggs, A. gossypii nymphs, and B. tabaci nymphs conforms to the Holling type II functional response model. Predation capacity was significantly affected by the predator’s developmental stage, sex, and prey type, with older nymphs and male adults demonstrating the highest predation potential on A. gossypii nymphs. However, actual predation impact in the field was strongly limited by prey population density, spatiotemporal availability, and regional ecological conditions. No effective predation on H. armigera eggs was documented, while predation on A. gossypii displayed clear regional heterogeneity. The acquisition of animal nutrition through facultative predation may improve the population fitness and damage potential of A. lucorum under resource-limited conditions, thereby complicating integrated pest management (IPM) approaches. Therefore, within an IPM framework, it is recommended to implement a management system based on dynamic monitoring and ecological risk assessment. This approach supports region-specific and temporally targeted precision control strategies, which would help avoid unintended enhancement of the pest’s ecological competitiveness during management interventions.

Author Contributions

Conceptualization, K.W. and L.W.; methodology, L.W., B.L. and K.W.; software, L.W.; validation, L.W. and B.L.; formal analysis, L.W.; investigation, L.W. and B.L.; resources, K.W.; data curation, L.W.; writing—original draft preparation, L.W.; writing—review and editing, all; visualization, L.W.; supervision, K.W.; project administration, K.W.; funding acquisition, K.W. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Academician Workstation of the Agricultural High-Tech Industrial Area of the Yellow River Delta, National Center of Technology Innovation for Comprehensive Utilization of Saline-Alkali Land. This funding source does not have a grant number.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding author.

Acknowledgments

We are grateful to the Plant Protection Institute of the Henan Academy of Agricultural Sciences for their assistance during the field investigation and for providing the test samples from Xinxiang.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Predation responses of Apolygus lucorum at different developmental stages to three prey species: Helicoverpa armigera eggs (AD), Aphis gossypii nymphs (EH), and Bemisia tabaci nymphs (IL). Each subfigure represents a specific developmental stage: 2nd instar nymphs (A,E,I), 4th instar nymphs (B,F,J), 5-day-old female adults (C,G,K), and 5-day-old male adults (D,H,L). Data are presented as mean daily predation rate ± SE (n = 5 per prey density). Curves were fitted using the Holling type II disc equation. Significant differences in prey consumption among developmental stages were analyzed using one-way ANOVA followed by Tukey’s HSD test. Colors in the figure represent different developmental stages: orange, 2nd instar nymph; green, 4th instar nymph; purple, 5-day-old female; blue, 5-day-old male.
Figure 1. Predation responses of Apolygus lucorum at different developmental stages to three prey species: Helicoverpa armigera eggs (AD), Aphis gossypii nymphs (EH), and Bemisia tabaci nymphs (IL). Each subfigure represents a specific developmental stage: 2nd instar nymphs (A,E,I), 4th instar nymphs (B,F,J), 5-day-old female adults (C,G,K), and 5-day-old male adults (D,H,L). Data are presented as mean daily predation rate ± SE (n = 5 per prey density). Curves were fitted using the Holling type II disc equation. Significant differences in prey consumption among developmental stages were analyzed using one-way ANOVA followed by Tukey’s HSD test. Colors in the figure represent different developmental stages: orange, 2nd instar nymph; green, 4th instar nymph; purple, 5-day-old female; blue, 5-day-old male.
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Figure 2. Population dynamics of Apolygus lucorum, Helicoverpa armigera eggs, and Aphis gossypii in cotton fields, and the positive detection rate of A. gossypii DNA in field-collected A. lucorum individuals. Data are shown for Langfang (A,B), Xiajin (C,D), and Xinxiang (E,F) in 2009 (A,C,E) and 2010 (B,D,F). Left y-axis indicates the number of A. lucorum adults and H. armigera eggs per 100 plants, as well as the positive detection rate (%) of A. gossypii DNA. Right y-axis indicates the number of A. gossypii nymphs per 100 plants. Data are presented as mean ± SE (n = 5 sampling points per date).
Figure 2. Population dynamics of Apolygus lucorum, Helicoverpa armigera eggs, and Aphis gossypii in cotton fields, and the positive detection rate of A. gossypii DNA in field-collected A. lucorum individuals. Data are shown for Langfang (A,B), Xiajin (C,D), and Xinxiang (E,F) in 2009 (A,C,E) and 2010 (B,D,F). Left y-axis indicates the number of A. lucorum adults and H. armigera eggs per 100 plants, as well as the positive detection rate (%) of A. gossypii DNA. Right y-axis indicates the number of A. gossypii nymphs per 100 plants. Data are presented as mean ± SE (n = 5 sampling points per date).
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Table 1. Logistic regression parameters and Holling Type II model goodness-of-fit for the functional responses of different developmental stages of Apolygus lucorum to three prey species.
Table 1. Logistic regression parameters and Holling Type II model goodness-of-fit for the functional responses of different developmental stages of Apolygus lucorum to three prey species.
PreyDevelopmental Stage ParameterEstimateStandard Errorp-ValueR2
Helicoverpa armigera eggs2nd instar nymphLinear (P1)−0.06480.09400.49060.9199
Quadratic (P2)0.00340.00350.3415
4th instar nymphLinear (P1)−0.62350.50080.21320.9736
Quadratic (P2)0.01600.01470.2743
5-day-old female adultLinear (P1)−0.67540.19240.00040.9784
Quadratic (P2)0.01980.00610.0012
5-day-old male adultLinear (P1)−0.11820.11040.28440.9965
Quadratic (P2)0.00290.00400.4744
Aphis gossypii nymph2nd instar nymphLinear (P1)−0.30190.36300.40560.9655
Quadratic (P2)0.01810.03360.5894
4th instar nymphLinear (P1)−0.31960.37350.39220.9802
Quadratic (P2)0.03560.03500.3086
5-day-old female adultLinear (P1)−0.18790.37280.61420.9637
Quadratic (P2)0.03500.03570.3246
5-day-old male adultLinear (P1)−0.35900.45130.42630.9573
Quadratic (P2)0.04510.04220.2858
Besimia tabaci nymph2nd instar nymphLinear (P1)−0.15970.36870.66490.9141
Quadratic (P2)0.01230.03590.7315
4th instar nymphLinear (P1)−0.10050.27030.71000.9540
Quadratic (P2)0.01080.02570.6750
5-day-old female adultLinear (P1)−0.28900.29940.33440.9029
Quadratic (P2)0.03310.02820.2404
5-day-old male adultLinear (P1)−0.05300.29840.85930.9556
Quadratic (P2)0.00250.02880.9320
Table 2. Functional response parameters of Apolygus lucorum at different developmental stages preying on eggs of Helicoverpa armigera, nymphs of Aphis gossypii, and nymphs of Bemisia tabaci.
Table 2. Functional response parameters of Apolygus lucorum at different developmental stages preying on eggs of Helicoverpa armigera, nymphs of Aphis gossypii, and nymphs of Bemisia tabaci.
PreyDevelopmental Stage Functional ResponsepaTh (d)Maximum Consumptiona/Th
Helicoverpa armigera eggs2nd instar nymphNa = N/(3.1989 + 0.0239 N)0.00100.31260.023941.8413.08
4th instar nymphNa = N/(0.9109 + 0.0145 N)0.00021.09780.014568.9775.71
5-day-old female adultNa = N/(0.9113 + 0.0176 N)<0.00011.09730.017656.8262.35
5-day-old male adultNa = N/(1.0055 + 0.0136 N)<0.00010.99450.013673.5373.13
Aphis gossypii nymph2nd instar nymphNa = N/(0.9358 + 0.1254 N)0.00481.06860.12547.978.52
4th instar nymphNa = N/(1.4069 + 0.0037 N)<0.00010.71080.0037270.27192.11
5-day-old female adultNa = N/(1.4640 + 0.0025 N)<0.00010.68310.0025400.00273.24
5-day-old male adultNa = N/(1.2265 + 0.0012 N)<0.00010.81530.0012833.33679.42
Besimia tabaci nymph2nd instar nymphNa = N/(5.9340 + 0.2886 N)<0.00010.16850.28863.470.58
4th instar nymphNa = N/(3.7406 + 0.0597 N)0.00030.26730.059716.754.48
5-day-old female adultNa = N/(5.2068 + 0.2926 N)0.02000.19210.29263.420.66
5-day-old male adultNa = N/(3.6654 + 0.1742 N)<0.00010.27280.17425.741.57
p values indicate the significance level of the Holling type II model fitting. a is the attack rate, and Th is the handling time (d). Maximum daily prey consumption was calculated as 1/Th, and predation capacity was calculated as a/Th. The total experimental time was T = 1 d. All treatments conformed to the Holling type II model based on functional response curve fitting.
Table 3. Positive detection rates of Helicoverpa armigera eggs and Aphis gossypii in cotton fields from three regions during 2009–2010.
Table 3. Positive detection rates of Helicoverpa armigera eggs and Aphis gossypii in cotton fields from three regions during 2009–2010.
YearAreaTotal No. of DetectionsPositive Detection Rate of Helicoverpa armigera EggsPositive Detection Rate of Aphis gossypii
2009Langfang1260.00 7.04 ± 3.73 a
Xiajin1410.00 1.22 ± 1.26 a
Xinxiang420.00 0.00 ± 2.47 a
2010Langfang1350.00 3.70 ± 2.47 a
Xiajin1140.00 0.00 ± 1.23 a
Xinxiang410.00 0.00 ± 1.23 a
Data for positive detection rate are presented as means ± SE. Means in the same column followed by different letters are significantly different for each year (p < 0.05) (one-way ANOVA, Tukey’s HSD).
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Wang, L.; Liu, B.; Wu, K. Evaluation of Predation on Phytophagous Insects by a Phytozoophagous Mirid Bug, Apolygus lucorum. Insects 2026, 17, 397. https://doi.org/10.3390/insects17040397

AMA Style

Wang L, Liu B, Wu K. Evaluation of Predation on Phytophagous Insects by a Phytozoophagous Mirid Bug, Apolygus lucorum. Insects. 2026; 17(4):397. https://doi.org/10.3390/insects17040397

Chicago/Turabian Style

Wang, Lili, Baoyou Liu, and Kongming Wu. 2026. "Evaluation of Predation on Phytophagous Insects by a Phytozoophagous Mirid Bug, Apolygus lucorum" Insects 17, no. 4: 397. https://doi.org/10.3390/insects17040397

APA Style

Wang, L., Liu, B., & Wu, K. (2026). Evaluation of Predation on Phytophagous Insects by a Phytozoophagous Mirid Bug, Apolygus lucorum. Insects, 17(4), 397. https://doi.org/10.3390/insects17040397

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