Methodology for rearing, development, and reproduction of Iridopsis panopla Prout (Lepidoptera: Geometridae) on different diets

Metodologia de criação, desenvolvimento e reprodução de Iridopsis panopla (Lepidoptera: Geometridae) em diferentes dietas

Abstract

Damage caused by insect pests, such as Iridopsis panopla Prout, 1932 (Lepidoptera: Geometridae), has been frequent and can reduce the volume of timber produced in eucalyptus plantations by up to 40%. This study aimed to establish a laboratory rearing methodology and evaluate the development and reproduction of Iridopsis panopla on different diets. The experiments were conducted at the Insect Biotechnology Laboratory (BIOINSECTH) of the Universidade Federal da Grande Dourados, Mato Grosso do Sul, under controlled conditions (25 ± 2 °C, 70 ± 10% RH, and a 14-hour photoperiod). We evaluated larval duration and viability, pre-pupa period and viability, morphological characteristics of caterpillars, pupae, and eggs, sexual dimorphism, as well as pre-oviposition, oviposition, and post-oviposition periods. The width of the cephalic capsule and larval length were measured daily from hatching to the prepupa stage. Five treatments were tested: a diet of Chrysodeixis includens combined with leaves from clone I144 (Eucalyptus urophylla × E. grandis) (D1); a diet of C. includens (D2); a diet of Spodoptera frugiperda combined with leaves from clone I144 (D3); a diet of S. frugiperda (D4); and isolated leaves of clone I144 (D5). The diet significantly influenced the biological performance of the species. The combination of leaves from clone I144 (Eucalyptus urophylla × E. grandis) with a C. includens diet (D1) resulted in the best development and reproduction. Diet D4 proved unviable, and the exclusive provision of leaves from clone I14. Iridopsis panopla completed five larval instars on viable diets, with variation in the interval between molts depending on the treatment. Iridopsis panopla can be successfully reared in the laboratory, showing better performance when fed with leaves from clone I144 combined with the artificial diet of Chrysodeixis includens.

Keywords:
Eucalyptus urograndis; defoliating caterpillar; pest; laboratory rearing

Resumo

Danos causados por insetos-praga, como Iridopsis panopla Prout, 1932 (Lepidoptera: Geometridae), têm sido frequentes e podem reduzir em até 40% o volume de madeira produzido em plantações de eucalipto. Este estudo objetivou estabelecer uma metodologia de criação em laboratório e avaliar o desenvolvimento e a reprodução de Iridopsis panopla em diferentes dietas. Os experimentos foram conduzidos no Laboratório de Biotecnologia de Insetos (BIOINSECTH) da Universidade Federal da Grande Dourados, Mato Grosso do Sul, em condições controladas (24 ± 2 °C, 70 ± 20% UR e fotoperíodo de 14 h). Avaliou-se a duração e viabilidade larval, período e viabilidade de pré-pupas, características morfológicas de lagartas, pupas e ovos, dimorfismo sexual, além dos períodos de pré-oviposição, oviposição e pós-oviposição. Mediu-se diariamente a largura da cápsula cefálica e o comprimento larval desde a eclosão até a pré-pupa. Foram testados cinco tratamentos: dieta de Chrysodeixis includens combinada com folhas do clone I144 (D1); dieta de C. includens (D2); dieta de Spodoptera frugiperda combinada com folhas do clone I144 (D3); dieta de S. frugiperda (D4); e folhas do clone I144 (Eucalyptus urophylla × E. grandis) isoladas (D5). A dieta influenciou significativamente o desempenho biológico da espécie. A combinação de folhas do clone I144 (Eucalyptus urophylla × E. grandis) com dieta de C. includens (D1) proporcionou o melhor desenvolvimento e reprodução. A dieta D4 mostrou-se inviável e o fornecimento exclusivo de folhas do clone I144 (D5) resultou em baixa sobrevivência. Iridopsis panopla completou cinco ínstares larvais nas dietas viáveis, com variação no intervalo entre mudas conforme o tratamento. Iridopsis panopla pode ser criada com sucesso em laboratório, apresentando melhor desempenho quando alimentada com folhas do clone I144 associadas à dieta artificial de Chrysodeixis includens.

Palavras-chave:
Eucalyptus urograndis; lagarta desfolhadora; praga; criação em laboratório

1. Introduction

In 2023, approximately 76% of Brazil’s forest plantation area was occupied by eucalyptus species, totaling approximately 7.6 million hectares (IBÁ, 2023). The eucalyptus production cycle in Brazil is the shortest in the world, ranging from five to seven years (Lopes et al., 2022). This characteristic helps reduce pressure on native areas, promotes the conservation of water and soil resources, and enables the sequestration of up to 184 tons of carbon dioxide per hectare by the end of the cycle (Cândido et al., 2014).

The monoculture model adopted in these plantations promotes the emergence of pest outbreaks, which occur more frequently and with greater intensity than in more diverse forest environments. When not properly managed, these outbreaks can cause significant economic losses (Guedes et al., 2000; Teixeira and Rodrigues, 2021).

Among the main groups of pests affecting eucalyptus, defoliating caterpillars stand out; they consume the leaf blade and, at high population densities, can completely defoliate the plants, compromising their growth (Lucchetta et al., 2022).

This group is considered the second most significant pest for the crop, second only to leaf-cutting ants (Guedes et al., 2000). In recent years, severe outbreaks of Iridopsis panopla Prout (Lepidoptera: Geometridae) have been recorded in eucalyptus-growing areas, particularly in the states of Mato Grosso do Sul and São Paulo, affecting more than 860,000 hectares in 2021 (Santos et al., 2024; Wilcken et al., 2022).

Despite the increasing occurrence of I. panopla, studies on biology, morphology, and feeding behavior remain scarce. To date, there are no records of rearing methods or appropriate diets for maintaining this species under controlled conditions. This information is essential for understanding its potential for damage and for informing the development of alternative management strategies.

Considering this gap, this study was conducted with the aim of establishing a rearing methodology and evaluating the growth and reproduction of I. panopla on different diets, including leaves from eucalyptus clone I144 (a hybrid of Eucalyptus urophylla × E. grandis) and artificial diets.

2. Materials and Methods

In June 2022, specimens of defoliating lepidoptera were collected from a commercial plantation of urograndis in Brasilândia, Mato Grosso do Sul. Adults were collected using an FTF-model cage (24 × 24 cm) made of nylon mesh. After collection, the material was taken to the Laboratory of Biological Control of Insects (LECOBIOL) (22° 11’56.56’’ S, 54° 56’ 1.396” W) at the Faculty of Biological and Environmental Sciences (FCBA) of the Universidade Federal da Grande Dourados, Mato Grosso do Sul, Brazil, where they were kept in a climate-controlled room at 25 ± 2 °C, 70 ± 10% RH, and a 14-hour photoperiod.

2.1. Species identification

Adult lepidopterans were collected in the field by Carlos Frederico Wilcken and Fábio Araújo dos Santos and sent to Dr. Alberto Soares Côrrea at the Laboratory of Molecular Ecology of Arthropods, Luiz de Queiroz College of Agriculture, University of São Paulo (ESALQ/USP), in Piracicaba, São Paulo, Brazil, for species identification using molecular techniques. Thirty-two adults underwent total genomic DNA extraction according to the methodology described by Ferronato et al. (2019). The mitochondrial cytochrome c oxidase subunit I (COI) gene region was amplified and sequenced. The resulting sequences were compared with those available in the BOLD Systems (2026) reference database, and showed 100% sequence similarity with I. panopla (Santos, 2023a).

2.2. Rearing methodology for Iridopsis panopla

To feed adult I. panopla specimens collected in the field, a 10% honey solution was used, soaked into cotton balls and placed on the upper outer part of the entomological cages. This solution served as an energy source during the reproductive period. The females laid eggs preferentially on the rough fabric of the cages, often between the fabric and the cotton balls, which facilitated egg collection.

The eggs remained in the cages until the caterpillars hatched. The newly hatched caterpillars were carefully removed with a brush, using the silk thread secreted by the species through its mouthparts as a handle for handling. They were then placed in plastic containers in groups of varying sizes, with the aim of observing initial adaptation and determining the optimal number of individuals per experimental unit.

The jars used had a capacity of 120 mL and contained leaves from eucalyptus clone I144 (a hybrid of E. urophylla × E. grandis) (Figure 1). The leaf petioles were inserted into plastic microtubes (1.5 mL Eppendorf tubes) filled with water, which were placed in an open hole in the jar lids to slow the dehydration of the plant material. The leaves and microtubes were replaced every other day to ensure the quality of the diet was maintained.

Figure 1
Setup of the laboratory rearing methodology for I. panopla. (1) Adult I. panopla; (2) Cage used for feeding and oviposition by adults; (3) Eppendorf tube with water to prevent dehydration of the eucalyptus leaves; (4) Artificial diet containing wood stick fragments to support the caterpillars; (5) Rearing system for early instars, with eucalyptus leaves and diet; (6) Bouquet of eucalyptus leaves in a Becker container, intended for larger instars; (7) Cage adapted for application of the bouquet technique; (8) Pupae collected from the bottom of the rearing cages, subsequently transferred to the adult cage. Dourados - MS, UFGD, 2026.

To ensure adequate ventilation and prevent the insects from escaping, 2-cm openings were made in the lids of the jars. Absorbent paper was inserted between the lid and the container to serve as an addition al seal. The artificial diets were provided in pieces approximately 2 cm in size, with wooden sticks placed at the ends to support the food and facilitate access for the caterpillars.

As the caterpillars reached about 2 cm in length, they were transferred to bundles of eucalyptus leaves, kept in larger entomological cages (40 cm long and deep, and 30 cm wide). This stage allowed for monitoring of their development up to the prepupal stage. The prepupae were collected from the bottom of the larger cages, placed in Petri dishes with sand and eucalyptus leaves to simulate field conditions, and transferred to smaller cages (15 cm in height, width, and depth), where they remained until the adults emerged.

The emerged adults were fed daily with the same 10% honey solution, applied to cotton swabs, ensuring the continuity of the reproductive cycle under controlled conditions. The entire process was conducted with technical rigor, aiming to standardize experimental conditions and obtain reliable data on the development and reproduction of the species under different diets.

2.3. Experimental procedure

Groups of six newly emerged second-generation caterpillars were separated into plastic pots representing a plot with seven replicates and fed according to the following treatments: a diet of Chrysodeixis includens (Walker, 1858) (Lepidoptera: Noctuidae) and urograndis clone I144 leaves (D1), a diet of C. includens (D2), a diet of Spodoptera frugiperda J. E. (Smith, 1797) (Lepidoptera: Noctuidae) and urograndis clone I144 leaves (D3), a diet of S. frugiperda (D4) and urograndis clone I144 leaves (D5) (Table 1). The number of caterpillars per replicate was determined in a pretest, during which larval development and viability under different densities per pot were observed. Diets D1 and D2 were developed for C. includens (Greene et al., 1976) and S. frugiperda (Nalin, 1991) caterpillars, respectively (Table 2).

Table 1
Feeding of I. panopla on leaves of urograndis clone I144 + artificial diets (D1 and D2), artificial diets (D1 and D2), and clone leaves alone. Dourados, MS, UFGD, 2026.

Treatment	Food
D1	Urograndis leaf, clone I144 + Diet 1 (D1)
D2	Diet 1 (D1)
D3	Urograndis leaf, clone I144 + Diet 2 (D2)
D4	Diet 2 (D2)
D5	Urograndis leaf

Table 2
Composition of diets formulated for C. includens larvae (Greene et al., 1976) (D1) and S. frugiperda larvae (D2) (Nalin, 1991), respectively. Dourados, MS, UFGD, 2026.

Ingredients	Quantities
	Diet 1 (D1)	Diet 2 (D2)
Wheat germ (g)	270	79.5
Yeast (g)	168.9	50.5
Soy protein (g)	137	*
Casein (g)	135	*
Ascorbic acid (g)	16.2	5.1
Sorbic acid (g)	8.1	1.65
Nipagin (g)	13.5	3.15
Tetracycline (g)	0.54	*
Agar (g)	105	20.5
Vitamin solution (mL)	42	*
Formaldehyde (mL)	16.2	1.0
Distilled water (L)	5.4	1.2
Beans (g)	377.5	165

• ^*
  Ingredients not used in this diet.

2.3.1. STAGE I - Biological and reproductive aspects of I. panopla on different diets

The caterpillars were monitored daily throughout the feeding period to determine larval duration (LD), larval viability (LV), the prepupal period (PPP) defined as the interval between the cessation of feeding and the transition to the pupal stage, and prepupal viability (PPV). In addition, morphological traits, including the color and markings of the caterpillars, the shape and coloration of the pupae, sexual dimorphism, and the texture, shape, and size of the eggs, were assessed using a magnifying glass.

Pupae of I. panopla were photographed, sexed, weighed, identified, and individually tagged, and their duration (DP), weight (PP), length (CP), viability (VP), and sex ratio (RS) were determined. Six cages, made from 30 cm lengths of 100 mm diameter PVC pipe with a rough fabric serving as an oviposition substrate on top and, above that, cotton soaked in a honey solution, were set up for each insect pair per treatment.

The remaining adults were kept in 120-mL jars and fed 10% honey on cotton placed in a hole in the jar lids to determine the lifespan of male and female specimens of this insect. The rough polypropylene fabric, used as an oviposition substrate, and the food were replaced daily, and the former was placed in labeled glass Petri dishes (15 cm × 2.5 cm) to count the eggs per day and assess their viability based on caterpillar hatching.

The pre-oviposition, oviposition, and post-oviposition periods of I. panopla in PVC cages were also evaluated. The pre-oviposition period corresponded to the interval between the emergence of adults and the start of oviposition, the oviposition period to the interval between the first and last egg-laying, and the post-oviposition period to the interval between the last egg-laying and the death of the adult.

2.4. Statistical analysis

Data on larval duration (DL) and viability (VL), pre-pupa period (PPP) and viability (VPP), pupal duration (DP), weight (PP), length (CP), pupal viability (VP), sex ratio (SR), and pre-oviposition, oviposition, and post-oviposition periods were subjected to analysis of variance, and means were compared using Tukey’s test at a 5% significance level. The experiment was conducted using a randomized block design, with five treatments and seven replicates per treatment. Each replicate consisted of a pot containing six larvae.

2.4.1. STAGE II - Development of I. panopla on different diets

The width of the cephalic capsule and the body length of three I. panopla larvae from the ongoing experiment were recorded daily from hatching through the prepupal stage, using images captured with a Zeiss Discovery.V8 magnifying glass and analyzed using ZEN 2.3 Lite software. The progression of instars was determined based on measurements of the cephalic capsule and body length. The change in instar was identified by the replacement of the cephalic capsule, a characteristic that marks the beginning of a new developmental stage.

2.5. Statistical analysis

Data on body growth and head capsule size were analyzed using nonparametric tests due to the longitudinal nature of the data and the presence of missing observations due to mortality. Differences between treatments were assessed using the Kruskal-Wallis test, followed by pairwise multiple comparisons using the Mann-Whitney U test.

Data on body growth and head capsule size were analyzed using nonparametric tests due to the longitudinal nature of the data and the presence of missing observations due to mortality. Differences between treatments were assessed using the Kruskal-Wallis test, followed by pairwise multiple comparisons using the Mann-Whitney U test.

All statistical analyses were performed using Python (version 3.12), with a significance level set at α = 0.05. Assumptions of normality were assessed using the Shapiro-Wilk test, and due to a violation of normality in the residuals, nonparametric methods were chosen. The growth curves include confidence intervals based on the standard error of the mean.

3. Results

3.1. STAGE I - Biological and reproductive aspects of I. panopla on different diets

The rearing methodology for I. panopla allowed the species life cycle to be maintained under controlled conditions. The set of procedures adopted ensured stable conditions for the development of the caterpillars and allowed for an accurate assessment of the effect of different diets on the species life cycle.

Under these conditions, the larval duration of I. panopla showed marked variation among the treatments: it was shortest in D1, intermediate in D3 and D5, and longest in D2. In treatment D4, the caterpillars remained alive for up to 65 days without advancing to the second instar, indicating an inability to develop on this diet (Table 3).

Table 3
Larval duration (LD) and viability (LV), pre-pupal period (PPP), pre-pupal duration (PD) (days), and viability (VPP) (%) (Means ± SE) of I. panopla fed diets of C. includens and leaves of clone I144 (a hybrid of E. urophyla × E. grandis) (D1), C. includens diet (D2), S. frugiperda diet and leaf of clone I144 (D3), S. frugiperda diet (D4), and leaf of clone I144 (D5). Dourados - MS, UFGD, 2026.

Diets	DL	VL	PPP	DP	VPP (%)
D1	37.06 ± 0.47a	59.51 ± 7.14a	2.20 ± 0.17a	14.09 ± 0.26a	79.29 ± 2.65b
D2	57.2 ± 2.44c	16.66 ± 0.00b	2.00 ± 0a	13.00 ± 0.58a	100.00 ± 0a
D3	43.65 ± 1.39b	64.28 ± 6.73a	2.52 ± 0.2a	13.33 ± 0.43a	100.00 ± 0a
D4	*	*	*	*	*
D5	47.07 ± 2.11b	28.47 ± 7.04b	2.53 ± 0.36a	13.2 ± 0.37a	100.00 ± 0a

• ^*
  Caterpillars did not pass the first instar. Means followed by the same letter in the column do not differ by Tukey's test (p≤0.05).

Only one I. panopla caterpillar survived in most plots of treatment D5. In treatment D2, the caterpillars showed initial growth but died during molting, especially in the prepupal stage. The bodies of the I. panopla caterpillars were initially greenish-yellow, and their heads were brown with spots, except for those fed only on eucalyptus leaves, which had a greener hue.

Iridopsis panopla caterpillars initially scraped the underside of the leaves; from the third instar onward, when the cephalic capsule reached a width of approximately 1.05 mm, they began feeding on the edges of the leaves. Changes in the caterpillars bodies were observed starting from this instar, beginning with black dorsal spots (Figure 2), which increased to as many as four as they developed further. They remain in specific positions where they camouflage themselves as twigs or assume an arched shape.

Figure 2
Stages of development of the I. panopla caterpillar reared in the laboratory. (1) I. panopla caterpillar in the 1st instar; (2) 2nd instar; (3) 3rd instar; (4) 4th instar; (5) Fifth instar; (6) Prepupa; (7) Pupa; (8) Adult pair, showing sexual dimorphism in the antennae; (9) Eggs. Federal University of Grande Dourados, Dourados, Mato Grosso do Sul, Brazil, 2022. Dourados - MS, UFGD, 2026.

The results obtained for the pre-pupal and pupal stages and for the viability of I. panopla demonstrate that diets D1, D2, D3, and D5 yielded similar performance, with mean values IN D1 of 2.20 days for the prepupal period, 14.09 days for the pupal period, and 79.29% viability (Table 3). The weight, length, viability, and sex ratio of I. panopla pupae were similar among diets D1, D2, D3, and D5, while no pupae were obtained on diet D4 (Table 4).

Table 4
Weight (PP), length (CP) and viability (VP) of pupae and sex ratio (RS) (Means ± SE) of I. panopla reared under different diets composed of treatments with diet of C. includens and leaves of clone I144 (hybrid of E. urophyla × E. grandis) (D1), diet of C. includens (D2), diet of S. frugiperda and leaves of clone I144 (D3), diet of S. frugiperda (D4) and leaves of clone I144 (D5). Dourados - MS, UFGD, 2026.

Diets	PP (g)	CP (mm)	VP (%)	RS
D1	0.06 ± 0a	11.66 ± 0.21a	85.71 ± 14.29a	0.49 ± 0.11a
D2	0.06 ± 0a	11.39 ± 0.26a	90.00 ± 10a	0.40 ± 0.24 a
D3	0.06 ± 0a	11.61 ± 0.22a	97.14 ± 2.86a	0.57 ± 0.1 a
D4	*	*	*	*
D5	0.07 ± 0a	11.59 ± 0.17a	100.00 ± 0a	0.28 ± 0.15 a

• ^*
  Insects did not develop. Means followed by the same letter in the column do not differ by Tukey's test (p≤0.05).

The pupae of I. panopla were brown in color, with the mesothoracic wing region green on the inside. The duration and viability of the pupae were similar across treatments. The pre-oviposition, oviposition, and post-oviposition periods of I. panopla did not differ between treatments D1, D3, and D5 (Table 5). Adults emerged and females oviposited on D2, but the number of pairs was insufficient for statistical testing.

Table 5
Pre-oviposition (PPP), oviposition (PO) and post-oviposition (PPO) periods (Means ± SE) in days; number of eggs (QO) and longevity of females (LF) and males (LM), in days, of I. panopla in treatments (Treat.) with diet of C. includens and leaves of clone I144 (hybrid of E. urophyla × E. grandis) (D1), diet of C. includens (D2), diet of S. frugiperda and leaves of clone I144 (D3), diet of S. frugiperda (D4) and leaves of clone I144 (D5). Dourados - MS, UFGD, 2026.

Trat.	PPP	PO	PPO	QO	LF	LM
D1	4.83±1.33a	12±2.39a	4.66±1.48a	196.66 ± 24.07a	15.5 ± 0.96a	16.85 ± 0.3a
D2	**	**	**	**	**	**
D3	7.00 ± 0.97 a	10.33 ± 2.11 a	4.16 ± 1.51 a	58.00 ± 11.92b	18.42 ± 1.08a	17.25 ± 1.83a
D4	*	*	*	*	*	*
D5	3.66 ± 0.33 a	11.33 ± 1.86 a	2 ± 0.58 a	58.00 ± 7.00b	14.5 ± 0.29a	15.87 ± 0.3a

• ^*
  Insects did not develop;
• ^**
  Insufficient number of couples. Means followed by the same letter in the column do not differ by Tukey's test (p≤0.05).

Adult I. panopla are light beige in color, with irregular horizontal brown markings; they have a cream-white abdomen with scattered dark gray scales and distinct sexual dimorphism, as evidenced by bipectinate antennae in males and setaceous antennae in females. The average lifespan of females was 15.5 ± 0.9 days and that of males was 16.85 ± 0.3 days (Table 5).

The number of eggs per female of I. panopla was higher on diet D1, with 196.66 (Table 5), showing a viability rate of 56.42%. Females laid eggs at night on the fabric located at the top of the PVC cage. The egg of I. panopla is flattened, with circular patterns on the chorion, light green in color, changing in hue to a darker pink until the caterpillar emerges. The incubation period was eight days.

3.2. STAGE II - Development of I. panopla on different diets

Iridopsis panopla exhibits five larval instars in the diets in which development occurred; the interval between head capsule molts varied depending on the diet provided. Throughout growth, the width of the cephalic capsule undergoes minor daily variations, but at the time of ecdysis there is a momentary reduction caused by internal pressure, followed by a marked expansion of the new capsule. In some cases, this expansion doubled the previous width (Figure 3), indicating a significant morphological reorganization between instars.

Figure 3
Growth curves for the cephalic capsule (mm) of I. panopla over time, comparing the five dietary treatments tested: urograndis leaves (clone I144), artificial diet D1, artificial diet D2, and combinations of leaves with D1 and D2. The curves represent the average growth per treatment, with shaded areas indicating confidence intervals. Dourados - MS, UFGD, 2026.

Treatment D1 (eucalyptus + Diet 1) showed the best overall performance, with accelerated growth of the cephalic capsule, reaching approximately 2.5 mm in 35-40 days, and the highest survival rate (70%). This result demonstrates a synergistic effect between eucalyptus and Diet 1, promoting both larval development and survival.

Treatment D4 (diet 2 alone) proved completely inadequate; the individuals remained in the second instar, with a maximum length of 6.35 mm (Figure 4), a cephalic capsule stunted at 0.5 mm, and 0% survival, with no progression to the prepupal stage. Treatment D5 (eucalyptus only) also showed limited performance, allowing only delayed and incomplete development, with 1% survival.

Figure 4
Body growth curves of I. panopla over approximately 65 days, comparing five dietary treatments: urograndis leaves (clone I144) D5, artificial diet D1, artificial diet D2, and combinations of leaves with D1 and D2. The lines represent the average growth per treatment, with shaded areas indicating the confidence intervals. Dourados - MS, UFGD, 2026.

Treatments D3 (eucalyptus + diet 2) and D2 (diet 1 alone) showed intermediate performance, both with a survival rate of 34%. However, D2 was less effective, with only 33% of the individuals reaching the prepupa stage, while T3 maintained more consistent growth, with the cephalic capsule reaching approximately 2.2 mm.

The high correlation between body length and cephalic capsule width (r = 0.97) confirms the robustness of the morphometric measurements and validates the use of the cephalic capsule as a reliable indicator of larval development, as it shows little variation within the instar. Furthermore, the criterion of a 2.2 cm head capsule size as a reference for entry into the prepupa stage was confirmed, contributing to the standardization of rearing protocols.

The survival curve (Figure 5) highlights the crucial role of eucalyptus as an essential nutritional component in larval development. The curves represent the duration of the larval stage until the transition to the prepupa; in treatment 4, the line indicates caterpillar mortality exclusively. Although diet D2 showed substantially reduced performance, characterized by a longer larval stage, and eucalyptus alone resulted in intermediate performance, the combination used in D1 demonstrated greater nutritional balance, reducing dependence on plant supplementation. Although D2 exhibited greater longevity (64 days), its development rate was lower, indicating that greater survival does not necessarily translate into better biological performance.

Figure 5
Survival curves of I. panopla caterpillars over time (days) across five dietary treatments: urograndis leaves (clone I144) D5, artificial diet D1, artificial diet D2, and combinations of leaves with D1 and D2, estimated using the Kaplan-Meier method. Dourados - MS, UFGD, 2026.

The results demonstrate that combining eucalyptus with specific artificial diets is essential for the proper development of I. panopla in the laboratory. Among the treatments evaluated, D1 stands out as the most effective for full development, while D3 represents the most balanced option for laboratory rearing protocols, as it combines high survival rates, consistent growth, and progression to the prepupal stage.

4. Discussion

4.1. STAGE I - Biological and reproductive aspects of I. panopla on different diets

The longer larval stage of I. panopla on diet D5, compared to diet D1, demonstrates that this diet may have provided a less balanced nutrient supply than the combination of leaves and artificial diet. Physical characteristics of the leaves, such as greater tissue toughness, leaf thickness, a more resistant cuticle, or the presence of trichomes, may have reduced feeding acceptance, ingestion, or nutrient assimilation and consequently prolonged larval development (Almeida et al., 2021).

The survival of only one caterpillar per replicate in D5 indicates that clone I144 leaves, when offered alone, were insufficient to sustain high larval survival under the laboratory conditions evaluated. Conversely, the improved performance observed when I144 leaves were combined with artificial diets suggests that this plant material has desirable characteristics for larval development, particularly when associated with a complementary nutritional source. The occurrence of field outbreaks further indicates that I. panopla can exploit this clone under natural conditions. Therefore, the low survival observed in D5 may be related to the conditions under which the plant material was produced and supplied, rather than being determined exclusively by the genetic characteristics of the host plant (Almeida et al., 2021).

The exclusive provision of either eucalyptus leaves or artificial diets may have contributed to the reduced survival of the caterpillars. The low viability of I. panopla on eucalyptus leaves may be due to clone I144, which has been reported as the least preferred by T. arnobia (Almeida et al., 2021). This preference may be chemical, physical, or morphological in nature, reducing food preference and, consequently, herbivory (Almeida et al., 2021).

The age of plants can have a significant effect on herbivores (Guedes et al., 2000). The use of leaves from six-month-old plants may have been inappropriate for I. panopla caterpillars, as outbreaks of defoliating insects of urograndis have been reported on plants in the intermediate stage, older than 65 months, as is the case with the species Sabulodes caberata Guenée, 1857 (Lepidoptera: Geometridae) (Andrade et al., 2016; Almeida et al., 2021). Older plants may favor the frequency and abundance of defoliating caterpillars due to the food richness that ensures the maintenance of larger populations, thereby providing greater carrying capacity (Guedes et al., 2000).

The initial scraping behavior observed in I. panopla caterpillars appears to be related to their small size in the early instars, which makes it difficult for them to cling to leaf edges. This pattern is consistent with that described for Leuciris fimbriaria (Stoll, 1781) (Lepidoptera: Geometridae), whose caterpillars also begin feeding by scraping the leaf surface before moving on to consume the margins (Heard et al., 2010). As they grow, the caterpillars of I. panopla begin to anchor themselves using their hind legs and feed on the leaf margins, a behavior similar to that of other Geometridae, reinforcing that feeding behavior is directly associated with body development.

In addition to its feeding behavior, I. panopla also exhibits defensive strategies comparable to those of L. fimbriaria. During the day, L. fimbriaria adopts postures that aid in camouflage, mimicking twigs or keeping its body arched. When disturbed, they release a silk thread and descend to the bottom of the cage, where they later pupate among plant debris (Heard et al., 2010). Iridopsis panopla exhibited similar behavior, suggesting that defense mechanisms and pupation site selection may be conserved within the Geometridae family. This similarity reinforces the idea that, despite differences in hosts and environments, closely related species tend to share adaptive behavioral strategies.

The similar results obtained for diets D1, D2, D3, and D5 regarding the prepupal and pupal stages and prepupal viability suggest that nutritional effects on I. panopla are predominantly manifested during the larval stage, while the impact tends to be reduced in subsequent stages. When compared to other Geometridae, the values obtained reinforce this interpretation. The prepupal period of I. panopla was similar to that of Nipteria panacea Thierry-Mieg, 1892 (Lepidoptera, Geometridae), which has an average of 1.9 days (Pratissoli et al., 2002).

However, the pupal duration was longer in I. panopla (14.09 days) than in N. panacea (11.6 days). This difference may reflect physiological characteristics of the species or the influence of the diets tested, suggesting that the pupal development of I. panopla may be more sensitive to nutritional variations than that of closely related species. Thus, although the pre-pupal period is consistent across species, the pupal stage appears to respond more variably to experimental conditions.

The absence of pupae in D4 indicates that the diet does not provide the minimum nutritional requirements for completing development, underscoring the importance of diet composition for larval survival. This result shows that, although I. panopla exhibits some dietary plasticity, its ability to complete its life cycle depends directly on the nutritional adequacy of the diet provided.

The similar pupal duration and viability of I. panopla across treatments indicate that, after completing larval development, the species maintains physiological stability regardless of the diet used. This pattern is consistent with that observed for I. ephyraria, which has a pupal period of 13 to 14.1 days when fed leaves from different natural hosts (Pinault and Quiring, 2008).

The similarity in the values suggests that species within the same genus tend to exhibit relatively similar pupal development, even when subjected to nutritional variations. However, the pupal weight of I. panopla (0.06 g) was lower than that recorded for T. arnobia fed on E. grandis at 25°C, which had a mean of 0.4143 g (Peres Filho and Berti Filho, 2003). This difference reinforces that, although pupal development time is similar among species, weight may vary, reflecting ecological and physiological differences between Geometridae and other families of Lepidoptera.

The pre-oviposition, oviposition, and post-oviposition periods of I. panopla, which were similar across treatments, confirm that the type of food on larval stages has no effect at this stage and highlight the importance of feeding adults a 10% honey solution. Nutrients at this stage affect parameters such as longevity and fecundity, with sugar being the primary energy source, composed of sucrose, glucose, and fructose (Zhou et al., 2023).

The coloration of I. panopla is similar to that of Iridopsis socoromaensis sp. n. (Lepidoptera: Geometridae), described as having wings with a mixture of whitish-gray and dark-gray scales, and a cream-white abdomen with scattered dark-gray scales (Vargas, 2021), a pattern also observed in I. panopla. The similar lifespan of males and females of I. panopla is consistent with that reported for N. panacea, with males living 19.5 days and females 19.4 days (Pratissoli et al., 2002).

The higher number of I. panopla eggs on diet D1 indicates greater adaptability of this diet, as reported for T. arnobia, which produced more eggs when its caterpillars were fed guava leaves than when fed E. grandis leaves. This may be due to the lower digestibility of eucalyptus leaves, as compounds such as tannins reduce protein availability for herbivores (Oliveira et al., 2005).

The number of eggs laid by I. panopla was similar to that of Macaria pallidata Packard (Warren, 1897) (Lepidoptera: Geometridae), with 182 eggs laid from the first night until the females death (Heard et al., 2010). Oviposition behavior without a specific frequency was also reported for N. panacea, with a similar number of eggs, 177 per female (Pratissoli et al., 2002).

Egg laying in the tissue may be related to the rough-textured substrate preferred by this insect for laying eggs in the field among the cracks in eucalyptus bark, similar to I. ephyraria laying eggs in cracks in tree bark (Pinault and Quiring, 2008).

The egg of I. panopla is flattened and bears circular patterns on the chorion; it is light green in color, changing to a darker pinkish hue until the caterpillar hatches, after which the chorion turns white. It is similar to that of I. ephyraria, and both species lay their eggs in cracks in tree bark (Pinault and Quiring, 2008). Variability in egg color during the embryonic period has also been reported for Iridopsis parrai Vargas (Lepidoptera: Geometridae), ranging from whitish green to pink (Vargas and Parra, 2013).

The habit of laying eggs in crevices or under the bark of host plants may be related to defense against biotic and abiotic factors, as reported for Epirrita autumnata (Borkhausen, 1794) (Lepidoptera: Geometridae), which lays eggs on a substrate covered by lichens to protect them from frost (Tammaru et al., 1995).

The results of this study demonstrate that the biological performance of I. panopla is directly influenced by the diet provided, with particular emphasis on the superiority of the diet combining urograndis leaves and a C. includens formulation (D1). The unsuitability of diet D4 and the low survival rate with the isolated I144 clone (D5) indicate nutritional limitations and possible plant resistance mechanisms. The feeding behavior, developmental pattern, and reproductive parameters observed reinforce the importance of balanced diets and the appropriate selection of plant hosts in rearing protocols. The results of this study contribute to the improvement of integrated management strategies and to the standardization of laboratory methodologies aimed at the study of I. panopla.

4.2. STAGE II - Development of I. panopla on different diets

Accurate determination of the number of instars is essential for studies in insect biology and for planning Integrated Pest Management strategies (Kalaimani et al., 2021). In the case of I. panopla, the identification of five instars in diets where development was successful is consistent with the pattern observed in other Geometridae. One example is Bupalus piniarius Linnaeus, 1758 (Lepidoptera: Geometridae), whose number of instars can range from four to eight under laboratory conditions, but which in the field generally completes its development in five or six stages, depending on environmental conditions (Gruys, 1970). The caterpillars of T. arnobia also exhibit similar values; after hatching, individuals of this species undergo six instars before pupating (Wilcken, 1996).

The width of the cephalic capsule proved to be a reliable morphometric parameter for distinguishing instars, as has been widely demonstrated in studies of Lepidoptera. The temporary reduction observed during ecdysis, followed by the rapid expansion of the new capsule, which in some cases doubled the previous width, reflects the physiological process of formation, hardening, and darkening of the newly synthesized cuticle (Chapman et al., 2013). Studies with T. arnobia confirm that analyzing the frequency distribution of cephalic capsule measurements across instars is a robust method for distinguishing larval stages, an approach that has also proven suitable for I. panopla (Wilcken, 1996).

The strong correlation between body length and cephalic capsule width indicates that both measurements are good indicators of the developmental stage. Nevertheless, the cephalic capsule offers an important methodological advantage: because it is a sclerotized structure, its dimensions remain stable within each instar, unlike body length, which varies according to the caterpillar’s feeding status and developmental stage (Wilcken, 1996). The experimental validation of the 2.2 cm value as a reference for entry into the prepupa stage contributes to the standardization of rearing protocols and facilitates monitoring of the life cycle under controlled conditions.

The poor performance observed in I. panopla on diet D4 is likely due to the diet’s insufficient nutritional composition or the absence of components essential for larval development. Evidence from other geometrid moths supports this interpretation; studies comparing the performance of T. arnobia on different hosts show that the species exhibits faster growth and greater developmental success on eucalyptus than on guava, indicating that chemical compounds or physical barriers present in the guava tree may negatively affect larval development (Holtz et al., 2003).

The results indicate that the contribution of eucalyptus leaves depended on the dietary context. The treatments combining leaves of clone I144 with artificial diets (D1 and D3) showed better biological performance than the exclusively artificial diets (D2 and D4), suggesting that the leaves provide nutritional components that were not fully supplied by the artificial formulations tested. However, the exclusive use of clone I144 leaves (D5) resulted in low survival, indicating that, although this plant material is a suitable host for the development of I. panopla, it may not provide all the nutrients required to maximize larval survival and development under the conditions evaluated. Therefore, the results support the use of clone I144 as a complementary food source, particularly in combination with the C. includens diet, rather than as a fully adequate diet when provided alone. Host-plant nutritional quality can directly affect larval duration, viability, and pupal weight, with consequences for adult reproductive performance, as previously demonstrated in studies of other species of the family Geometridae (Lemos et al., 1999).

The unsuitability of the D4 treatment and the unsatisfactory performance of the D2 treatment indicate that exclusive artificial diets, without supplementation with plant material, are not suitable for I. panopla in the formulations currently tested. The superiority of diets supplemented with eucalyptus reinforces this interpretation and is consistent with recent findings that identified I. panopla as one of the most common and abundant defoliating lepidopteran species in commercial eucalyptus plantations in the South and Southeast regions of Brazil, with population peaks between April and June (Santos, 2023b).

The results obtained in this study demonstrate that the larval development of I. panopla is strongly influenced by the diet provided, with the leaves of urograndis, clone I144, being an essential component for the proper progression of the life cycle. The precise identification of instars based on the cephalic capsule and body length reinforces the feasibility of standardized protocols for laboratory rearing.

The D4 diet proved to be inadequate, limiting growth during the second instar, while treatments that included eucalyptus (D1, D3, and D5) showed superior performance in terms of development and viability. Furthermore, the data show that longevity alone does not guarantee biological success, highlighting the importance of diets that promote the transition between life stages.

Acknowledgements

The Coordination for the Improvement of Higher Education Personnel (Capes) for awarding the scholarship. The National Council for Scientific and Technological Development (CNPq) for the funds provided to carry out this study. Association of Producers and Consumers of Planted Forests of Mato Grosso do Sul (REFLORE-MS) for its institutional support.

Data Availability Statement

Research data is only available upon request.

References

• ALMEIDA, C.A.C., GONÇALVES, F.S., RODRIGUES, M.B., SANTOS, J.M. and BREDA, M.O., 2021. Feeding preferences of Thyrinteina arnobia (Stoll, 1782) (Lepidoptera: Geometridae) on native and exotic hosts. Revista Árvore, vol. 45, no. 11, e4511. https://doi.org/10.1590/1806-908820210000011
  » https://doi.org/10.1590/1806-908820210000011
• ANDRADE, J.G.F., SÁ, V.G.M., LODI, S. and GODOY, B.S., 2016. Age of Eucalyptus urograndis plantations and occurrence of pest insects. Revista Árvore, vol. 40, no. 5, pp. 885-892. https://doi.org/10.1590/0100-67622016000500012
  » https://doi.org/10.1590/0100-67622016000500012
• BOLD SYSTEMS [online], 2026 [viewed 5 May 2026]. Available from: https://www.boldsystems.org/index.php/databases
  » https://www.boldsystems.org/index.php/databases
• CÂNDIDO, B.M., SILVA, M.L.N., CURI, N. and BATISTA, P.V.G., 2014. Post-planting water erosion in eucalyptus forests in the Paraná River basin, in eastern Mato Grosso do Sul. Brazilian Journal of Soil Science, vol. 38, no. 5, pp. 1565-1575. https://doi.org/10.1590/S0100-06832014000500022
  » https://doi.org/10.1590/S0100-06832014000500022
• CHAPMAN, R.F., SIMPSON, S.J. and DOUGLAS, A.E., 2013. The insects: structure and function 5th ed. Cambridge: Cambridge University Press, 929 p.
• FERRONATO, P., WOCH, A.L., SOARES, P.L., BERNARDI, D., BOTTON, M., ANDREAZZA, F. and CORRÊA, A.S., 2019. A phylogeographic approach to the Drosophila suzukii (Diptera: Drosophilidae) invasion in Brazil. Journal of Economic Entomology, vol. 112, no. 1, pp. 425-433. https://doi.org/10.1093/jee/toy321 PMid:30383249.
  » https://doi.org/10.1093/jee/toy321
• GREENE, G.L., LEPPLA, N.C. and DICKERSON, W.A., 1976. Velvetbean caterpillar: a rearing procedure and artificial medium. Journal of Economic Entomology, vol. 69, no. 4, pp. 487-488. https://doi.org/10.1093/jee/69.4.487
  » https://doi.org/10.1093/jee/69.4.487
• GRUYS, P., 1970. Growth in Bupalus piniarius (Lepidoptera: Geometridae) in relation to larval population density: 1. The influence of some abiotic factors on growth: 2. The effect of larval density. Wageningen: Wageningen University. PhD Thesis in Zoology. https://doi.org/10.18174/189345
  » https://doi.org/10.18174/189345
• GUEDES, R.N.C., ZANUNCIO, T.V., ZANUNCIO, J.C. and MEDEIROS, A.G.B., 2000. Species richness and fluctuation of defoliator Lepidoptera populations in Brazilian plantations of Eucalyptus grandis as affected by plant age and weather factors. Forest Ecology and Management, vol. 137, no. 1-3, pp. 179-184. https://doi.org/10.1016/S0378-1127(99)00326-6
  » https://doi.org/10.1016/S0378-1127(99)00326-6
• HEARD, T.A., ELLIOTT, L.P., ANDERSON, B., WHITE, L., BURROWS, N., MIRA, A., ZONNEVELD, R., FICHERA, G., CHAN, R. and SEGURA, R., 2010. Biology, host specificity, release and establishment of Macaria pallidata and Leuciris fimbriaria (Lepidoptera: Geometridae), biological control agents of the weed Mimosa pigra. Biological Control, vol. 55, no. 3, pp. 248-255. https://doi.org/10.1016/j.biocontrol.2010.08.001
  » https://doi.org/10.1016/j.biocontrol.2010.08.001
• HOLTZ, A.M., DE OLIVEIRA, H.G., PALLINI, A., MARINHO, J.S., ZANUNCIO, J.C. and OLIVEIRA, C.L., 2003. Adaptation of Thyrinteina arnobia to a new host and herbivore induced defense in eucalyptus. Pesquisa Agropecuária Brasileira, vol. 38, no. 4, pp. 453-458. https://doi.org/10.1590/S0100-204X2003000400002
  » https://doi.org/10.1590/S0100-204X2003000400002
• INDÚSTRIA BRASILEIRA DE ÁRVORES – IBÁ, 2023. Relatório anual 2023. São Paulo: IBÁ.
• KALAIMANI, T., NIRMALA, R., SWAMIAPPAN, M., SHANTHI, A., CHITRA, N. and VISWANATHAN, R., 2021. Instar determination for sugarcane internode borer Chilo sacchariphagus indicus (Kapur) (Lepidoptera: Crambidae). Journal of Asia-Pacific Entomology, vol. 24, no. 1, pp. 1087-1095. https://doi.org/10.1016/j.aspen.2020.08.010
  » https://doi.org/10.1016/j.aspen.2020.08.010
• LEMOS, R.N.S., CROCOMO, W.B., FORTI, L.C. and WILCKEN, C.F., 1999. Food selectivity and the influence of leaf age of Eucalyptus spp. on Thyrinteina arnobia (Lepidoptera: geometridae). Pesquisa Agropecuária Brasileira, vol. 34, no. 1, pp. 7-10. https://doi.org/10.1590/S0100-204X1999000100002
  » https://doi.org/10.1590/S0100-204X1999000100002
• LOPES, J.S., KIPERSTOK, A., FIGUEIRÊDO, M.C.B., ALMEIDA NETO, J.A. and RODRIGUES, L.B., 2022. Assessing the economic and environmental performance of cleaner production practices in eucalyptus planted forests using life cycle assessment. Journal of Cleaner Production, vol. 380, no. 2, pp. 134757. https://doi.org/10.1016/j.jclepro.2022.134757
  » https://doi.org/10.1016/j.jclepro.2022.134757
• LUCCHETTA, J.T., PEREIRA, F.F., CARDOSO, C.R.G., WILCKEN, C.F., SANTOS, F.A., PALOMBO, I.D., SANTOS, J.P., RAMOS, L.F.N., SIMONELI, B.C. and TAVARES, M.T., 2022. First record in Brazil of Brachymeria annulata (Fabricius) (Hymenoptera: Chalcididae) and Trichospilus diatraeae Cherian and Margabandhu (Hymenoptera: Eulophidae) parasitising pupae of Iridopsis panopla Prout (Lepidoptera: Geometridae) in eucalyptus (Myrtaceae). Canadian Entomologist, vol. 154, no. 1, e23. https://doi.org/10.4039/tce.2022.10
  » https://doi.org/10.4039/tce.2022.10
• NALIN, D.N., 1991. Biology, quantitative nutrition, and quality control of populations of Spodoptera frugiperda (J.E. Smith, 1797) (Lepidoptera: Noctuidae) on two artificial diets. Piracicaba: Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo.
• OLIVEIRA, H.N.D., ZANUNCIO, J.C., PEDRUZZI, E.P. and ESPINDULA, M.C., 2005. Rearing of Thyrinteina arnobia (Lepidoptera: Geometridae) on guava and eucalyptus in laboratory. Brazilian Archives of Biology and Technology, vol. 48, no. 5, pp. 801-806. https://doi.org/10.1590/S1516-89132005000600016
  » https://doi.org/10.1590/S1516-89132005000600016
• PERES FILHO, O. and BERTI FILHO, E., 2003. Thermal requirements of Thyrinteina arnobia (Stoll, 1782) and the effects of temperature on its biology. Ciência Florestal, vol. 13, no. 2, pp. 143-151. https://doi.org/10.5902/198050981750
  » https://doi.org/10.5902/198050981750
• PINAULT, L.L. and QUIRING, D.T., 2008. Sampling strategies and density-defoliation relationships for the pale-winged gray moth, Iridopsis ephyraria, on mature eastern hemlock. Forest Ecology and Management, vol. 255, no. 7, pp. 2829-2834. https://doi.org/10.1016/j.foreco.2008.01.058
  » https://doi.org/10.1016/j.foreco.2008.01.058
• PRATISSOLI, D., ZANUNCIO, J.C., OLIVEIRA, H.N. and FALQUETO, A., 2002. Biological characteristics of Nipteria panacea Thierry-Mieg (Lepidoptera, Geometridae), the avocado defoliator, in the mountainous region of Espírito Santo, Brazil. Revista Brasileira de Entomologia, vol. 46, no. 3, pp. 429-430. https://doi.org/10.1590/S0085-56262002000300008
  » https://doi.org/10.1590/S0085-56262002000300008
• SANTOS, F.A., 2023a. Biologia e manejo integrado da nova praga florestal lagarta-medideira-do-eucalipto Iridopsis panopla (Lepidoptera: Geometridae). Botucatu: Universidade Estadual Paulista.
• SANTOS, T.L., 2023b [viewed 11 February 2026]. Análise faunística de lepidópteros desfolhadores em plantios de eucalipto em três regiões do Brasil [online]. Botucatu: Universidade Estadual Paulista. Dissertação de Mestrado em Ciências Florestais. Available from: https://hdl.handle.net/11449/255334
  » https://hdl.handle.net/11449/255334
• SANTOS, F.H.M., PEREIRA, F.F., CARDOSO, C.R.G., LUCCHETTA, J.T., SANTOS, J.P., RAMOS, L.F.N., DIAS, L.S., ARAÚJO, H.M., SILVA, N.N.P., WILCKEN, C.F. and SANTOS, F.A., 2024. First record of Trichogramma pretiosum parasitizing Iridopsis panopla eggs in eucalyptus in Brazil. Fragmenta Entomologica, vol. 56, no. 2, pp. 20240042. https://doi.org/10.1515/flaent-2024-0042
  » https://doi.org/10.1515/flaent-2024-0042
• TAMMARU, T., KAITANIEMI, P. and RUOHOMÄKI, K., 1995. Oviposition choices of Epirrita autumnata (Lepidoptera: Geometridae) in relation to its eruptive population dynamics. Oikos, vol. 74, no. 2, pp. 296-304. https://doi.org/10.2307/3545659
  » https://doi.org/10.2307/3545659
• TEIXEIRA, G. and RODRIGUES, G.S.S.C., 2021. The geographical trajectory of forestry in Minas Gerais. Mercator, vol. 20, no. 4, e20004.
• VARGAS, H.A., 2021. Iridopsis socoromaensis sp. n., a geometrid moth (Lepidoptera, Geometridae) from the Andes of northern Chile. Biodiversity Data Journal, vol. 9, no. 1, e61592. https://doi.org/10.3897/BDJ.9.e61592 PMid:33551656.
  » https://doi.org/10.3897/BDJ.9.e61592
• VARGAS, H.A. and PARRA, L.E., 2013. Immature stages of Iridopsis parrai (Lepidoptera: Geometridae). Revista Colombiana de Entomología, vol. 39, no. 1, pp. 105-112. https://doi.org/10.1177/012004881303900118
  » https://doi.org/10.1177/012004881303900118
• WILCKEN, C.F., 1996. Biologia de Thrynteina arnobia (Stoll, 1782) (Lepidoptera: Geometridae) EM Espécies DE Eucalyptus e em dieta artificial. Piracicaba: Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo, 129 f. Tese de Doutorado em Entomologia.
• WILCKEN, C.F., SANTOS, F.A., BARBOSA, L.R., SOUZA, C.D. and ZANUNCIO, J.C., 2022. Desafios do manejo de lagartas desfolhadoras de eucalipto. Colombo: Embrapa Florestas.
• ZHOU, Y., SONG, P., LUO, D., FENG, Z., WANG, D. and HE, Y., 2023. Potential influence of nutrients intake by adults on the population dynamics of Athetis lepigone (Lepidoptera: Noctuidae). Journal of Asia-Pacific Entomology, vol. 26, no. 2, pp. 102104. https://doi.org/10.1016/j.aspen.2023.102104
  » https://doi.org/10.1016/j.aspen.2023.102104

*

e-mail: [e-mail]

Editor:

Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  05 Oct 2026
• Date of issue
  2026

History

• Received
  05 May 2026
• Accepted
  21 Aug 2026