Introduction

The Spanish cedar Cedrela odorata (L.) is one of the most economically and ecologically important tree species of the Meliaceae family in the tropical regions of America due to its productivity, high commercial timber value, and broad natural range (Cavers et al. 2004; Larrea et al. 2008; González-Luna and Cruz-Castillo 2021). Habitat fragmentation and illegal logging of C. odorata natural populations have reduced its adaptive and productive potential. Therefore, it has been recognized as a threatened species and has been included in the lists of international organizations such as International Union for Conservation of Nature (IUCN) and Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). It is also protected by NOM-059 in Mexico (SEMARNAT 2010; IUCN 2017; CITES 2021).

The establishment and productivity of commercial cedar plantations is compromised by attacks of the mahogany shoot borer, Hypsipyla grandella (Zeller) (Macías-Sámano 2001; Plath et al. 2011). Mated female shoot borers deposit their eggs on tree leaves (Griffiths 1997). Newly hatched larvae bore into the terminal shoot and create tunnels in the soft stem that ultimately kill the shoot. The plant will send out new shoots, but the tree will be forked or branched and its commercial value will be reduced (Briceño-Vergara 1997; Floyd and Hauxwell 2001; Lunz et al. 2010).

Chemical (Goulet et al. 2005), biological (Pulgarín et al. 2018), and silvicultural (Ruiz et al. 2016) control methods have been tested against this pest. Aside from being harmful to the environment, the application of conventional pesticides has also proven to be inefficient and economically unsustainable for various reasons, such as: larvae are inaccessible to pesticides since they are concealed in the stem, heavy rainfall washes away the applied pesticides, and a wide pesticide application window is required to protect the plants (Allan et al. 1976; Wylie 2001; Mahroof et al. 2002; Goulet et al. 2005). Therefore, the use of other control strategies as semiochemicals (kairomones and sex feromones) is recommended (Wyatt 1997). There are several advances in the study of semiochemicals with H. grandella, ranging from the toxic and repellent effects of extracts and oils obtained from different species of meliaceous, their ecological interactions with the volatiles of their host plant, and the use of sexual pheromone to control this pest (Holsten and Gara 1977; Borek et al. 1991; Soares 2002; Lago et al. 2006; Pineda-Ríos et al. 2016; Blassioli-Moraes et al. 2017; Borges et al. 2022).

The recognition of a host plant by insects is crucial to guarantee their survival and that of their progeny, since they not only perceive the plant as a food substrate, but also as appropriate sites for oviposition and refuge (Schoonhoven et al. 2005; Rojas 2012). Host plant recognition in insects is mainly mediated by blends of ubiquitous compounds, but also by specific compounds (Bruce et al. 2005). Insects are receptive to visual, olfactory, and gustatory cues during this process (Bernays and Champman 1994). Olfactory perception is involved in behavioral patterns associated with the search and selection of food sources (Libert et al. 2007), in the detection of toxic substances (Fuyama 1976), and in the recognition and selection of a partner (Billeter et al. 2009) and oviposition site (Hoffmann and O’Donnell 1990; Jaenike 1990). However, relatively little is known about the ecology, ethology, and biological interactions between H. grandella and C. odorata.

The susceptibility of C. odorata and Swietenia macrophylla (King) (monocultures as well as mixed plantations) to H. grandella attack has been documented, and the genetic variability of C. odorata has been studied (Newton et al. 1998; Cornelius and Watt 2003; Pérez-Salicrup and Esquivel 2008). These variations have implications in the production of attractants or deterrents for ovipositing H. grandella females (Honda 1995), and in morphology and branch formation (Grijpma 1976). To understand these phenomena, it is important to characterize and identify the compounds that mediate plant–insect interactions. Studies focussing on the relation between the moths H. grandella and Hypsipyla robusta (Moore) and different Meliaceae species have demonstrated the involvement of volatile compounds (most of the identified compounds are terpene derivatives, sesquiterpenes, and monoterpenes). These compounds elicit antennal responses and might be responsible for the attraction of these lepidopteran species (Soares et al. 2003; Lago et al. 2006; Abraham et al. 2014; Borges et al. 2022). Although these studies did not conduct behavior assays, they are a strong indication that H grandella uses volatiles from Meliaceae to locate the host plant. However, whether H. grandella is attracted by the volatiles emitted by cedar plants is still unknown.

In this study, we describe the nocturnal behavior and evaluate the attraction of H. grandella to cedar plants in field cages. We identify the volatile compounds emitted by healthy C. odorata plants and evaluate headspace volatiles from cedar plants and a synthetic blend in a Y-glass tube olfactometer.

Materials and Methods

Insects

First- to fourth-instar H. grandella larvae were collected from shoots and branches of five-year-old cedar trees. Sampling was conducted in April 2020 at the Unidad de Manejo para la Conservación de Vida Silvestre (UMA; registration ID: SEMARNAT-UMA-IN-1009-CHIS/17), located at the plantation “El Tesoro” (14°44′42.7" N, 92°14′37.2" W; 23.6 m a.s.l.) in Cantón Santa Lucía, municipality of Frontera Hidalgo, Chiapas, Mexico. The collected larvae were transported to the insectarium of the Laboratorio de Salud Forestal of El Colegio de la Frontera Sur (ECOSUR) in Tapachula, Chiapas, Mexico. They were reared on C. odorata shoots and maintained under the following controlled conditions: 25 ± 2 °C, 70 ± 20% relative humidity (RH), and a photoperiod of 12:12 h (L:D) during larval development (Vargas et al. 2001; Taveras et al. 2004). Pupae were sexed by the morphology of the genital opening (Sharma and Singh 1980). They were separated by sex and transferred to transparent plastic containers (250 ml). Emerging adults were then transferred to mating chambers (20 × 20 × 30 cm) in a 1:1 male:female ratio. Individuals were marked on the dorsal thorax with a Sharpie pen to differentiate females from males (Newell Brands, USA). Copulation was observed during the observation period (0:00–2:00 h), and its success was confirmed by the oviposition of fertile eggs (red color). Virgin female and male moths (1–2 d old) as well as mated females (3–4 d old) were selected for the experiments.

Plants

Cedrela odorata plants were obtained from the Santa Fe commercial plant nursery located in the municipality of Tuxtla Chico, Chiapas, Mexico. The plants were transplanted to containers (13 cm high, 12 cm diameter) filled with local soil, continuously irrigated and fertilized every 15 days with a nutrient solution (Steiner 1961). The plants were maintained under natural environmental conditions and covered with tulle netting to prevent insect damage. All the plants used in the study were one year old, with a heigh of 73 ± 2 cm and grown without pesticides.

Chemicals

The standards α-pinene, (E)-β-ocimene, 2-ethyl-1-hexanol, limonene, nonanal, and β-caryophyllene were purchased from Sigma-Aldrich (Toluca, Mexico) and were of 97–99% chemical purity according to the manufacturer. (E)-4,8-dimethyl-1,3,7-nonatriene (≥ 97%) was obtained from Pherobank BV (Wijk bij Duurstede, The Netherlands) and α-copaene (≥ 95%) from Cayman Chemicals (Ann Harbor, USA).

Bioassays

Behavior of Hypsipyla grandella. To determine the behavior of the insect towards the plant, a C. odorata plant was placed in the center of a cylinder-shaped field cage with a 3 m diameter and 2 m height (made from anti-aphid netting, mesh 40 × 25, cal. 0.009). One female (virgin or mated) was released into each cage for a period of 12 h (18:00–06:00 h), and insect activity observations were recorded every 10 min. The following behaviors were noted: movement of antennae, flight, walking, short (< 10 s) and long (> 30 s) wing fanning, calling position and oviposition. The observations were made with artificial red light (Coast-FL13) at a temperature of 22–28 ºC, 50–90% RH, and 0.10 ± 0.09 lx of ambient light. In total, 10 replicates were performed for each insect category.

Attraction to Cedrela odorata Plants. Cylinder-shaped field cages (diameter 3 m, height 2 m) were used to determine the attraction of H. grandella to the volatiles emitted by C. odorata. The test consisted of placing a cedar plant and a dummy control (cedar plant replica made from polyethylene, S1) within the cage separated 1 m from each other as a stimulus. A H. grandella moth (either virgin male, virgin female, or mated female) was released at a 1 m distance from the stimulus source. The moth was observed for 5 min and its first choice for the stimuli was recorded. These experiments were carried out between 19:00–23:00 h. The observations were made with artificial red light (Coast-FL13) at 25 ± 2 °C, 80 ± 10% RH and 0.03 ± 0.01 lx of ambient light. Moths, cedar plants, and dummy controls were used only once. In total, 50 replicates were performed for each insect category.

Attraction to Cedrela odorata Volatiles and Synthetic Blend. A Y-tube olfactometer (borosilicate glass, stem 15.52 cm; arms 12 cm, angled 45°, 2.5 cm i.d.) was used to evaluate the attraction of H. grandella towards C. odorata volatiles and a synthetic blend; 5 μl of volatile extract or 1 μl synthetic blend (stimulus) was loaded on a piece of Whatman No. 1 filter paper of 0.25 cm2 (Whatman International Ltd., Maidstone, UK). The stimulus was placed in one of the sample chambers, and 5 or 1 μl of dichloromethane (control) was placed in the second sample chamber. The synthetics blend contained 14.0 μg/μl α-pinene, 30.0 μg/μl 2-ethyl-1-hexanol, 6.0 μg/μl limonene, 3.0 μg/μl (E)-β-ocimene, 28.0 μg/μl nonanal, 18.0 μg/μl (E)-4,8-dimethyl-1,3,7-nonatriene, 21.0 μg/μl α-copaene and 5.0 μg/μl β-caryophyllene. The blend was made using a fivefold of the compounds found in the volatile emission of the plant (Table 1). The blend was prepared using dichloromethane as solvent. Activated charcoal filtered air at a rate of 0.5 l/min was pumped into each sample chamber. The flow was regulated with a pair of flowmeters and the air was humidified by passing it through a water jar placed before the olfactometer. Virgin males and females and mated females were tested in the bioassay. The moths were placed individually in the base of the stem of the Y-tube olfactometer. The first choice of the insect was registered if it crossed either of the two arms of the olfactometer (6 cm after intersection) within a 5 min period. The bioassay was stopped if the insect did not choose one of two samples within 5 min, in which case the moth’s response was recorded as non-responding. After each trial, the olfactometer was washed with distilled water and neutral soap, and dried in the oven at 120 ºC for 2 h. At the end of each bioassay, the position of the stimulus was changed to avoid experimental bias. The bioassays were conducted in a dark room between 19:00–23:00 h, and observation was facilitated by a red light (Coast-FL13) placed at 120 cm (0.4 lx) so the insects would not be disturbed (25 ± 1 °C and 65% RH). In total, 50 replicates were performed for each insect category.

Table 1 Average amount (ng/h/plant ± SE) of Cedrela odorata volatiles identified by GC–MS. Samples were collected over a 15 h scotophase period using a dynamic volatile collection system (N = 10)
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Volatile Collection

Volatile compounds were collected by using the dynamic headspace technique. Leaves and stems of a C. odorata plant were carefully enclosed within a 48 × 59 cm nylon oven bag (Reynolds, Lake Forest, USA). Volatiles were collected by passing air previously purified by an activated charcoal filter at a flow rate of 1 l/min with a vacuum pump (Model L-79200–00, Cole-Parmer, IL, United States). The volatiles were captured using Super Q adsorbent (50–80 mesh, 30 mg; ARS, Gainesville, USA) over a 15 h period (17:00–08:00 h) to capture the volatiles released mainly during the scotophase. At the end of each capture period, volatiles were eluted from the adsorbent with 400 μl of dichloromethane (HPLC-grade; Sigma-Aldrich, St. Louis, USA). The collected eluate was then concentrated to a final volume of 100 μl under a gentle stream of N2 and stored in small vials of 2 ml at -20 °C until analysis. A total of 10 cedar plants were used. However, since the plants released only small amounts of volatiles, it was necessary to repeat the extraction process ten times per each plant. The combined repetitions constituted the sample used for analysis. We evaluated the volatile GC–MS profiles of 10 samples. The collection of volatiles was performed at a temperature of 22–28 ºC and 80 ± 10% RH.

Identification of Volatiles

The volatile compounds from the headspace of cedar plants were identified with a Shimadzu GCMS-TQ8040 system (Shimadzu, Kyoto, Japan), composed of a TQ-8040 triple quadrupole mass spectrometer interfaced with a GC-2010 Plus gas chromatograph. A split-splitless capillary inlet system and a DB5-MS capillary column (30 m × 0.25 mm i.d.) were utilized for the analyses. All samples were injected in the splitless mode using the following temperature conditions: initial temperature of 50 ºC for 3 min, afterwards the temperature was increased by 15 °C/min to 280 ºC, and kept at a maximum of 280 °C for 10 min. Helium was utilized as the carrier gas at a constant flow rate of 1 ml/min. The injector port was set at 250 ºC. Ionization was via 70 eV electron ionization with an ion source temperature of 250 ºC. The software GC/MS Solution (version 4.20) was used for data processing. Each compound was identified preliminarily using the library of the National Institute of Standards and Technology (NIST, version 2014). The retention times and mass spectra of the identified compounds were compared with those of available synthetic standards to confirm their identification, and the retention index of each compound was determined. The concentrations of the identified compounds were determined by the external standard method and the release rate of each compound per plant per day was calculated, S2 (Sufang et al. 2013).

Statistical Analysis

The responses of H. grandella were analyzed by the G test with Williams’ correction in the R project software package (version 4.0.5; R Core Team 2021). Moths that did not choose either arm of the olfactometer within the 5 min observation period were excluded from analysis.

Results

Behavior of Hypsipyla grandella. Virgin and mated females spent most of their time resting (60%). The first recorded activity was the movement of antennae (13.5%), which began at 18:40 h, reached its activity peak at 19:00–23:30 h, and was registered for the last time one hour before the end of the experiment. The first and last recorded flight activities (8.5%) occurred at 18:50 h and 4:00 h, respectively, and peaked at 19:50–22:00 h. Walking activity (5.5%) was observed for the first time at 19:50 h and reached its maximum between 21:00 h and 22:50 h, after which sporadic activity bursts were registered until 2 h before the end of the experiment. Short wing-fanning (3%) was recorded for the first and last time at 19:00 h and 3:00 h, respectively. Maximum short wing-fanning activity was observed at 20:00–21:00 h, whereas long wing-fanning peaked (3%) between 3:00 h and 4:00 h. Virgin females assumed a calling position (5%) (by bending the abdomen dorsally up between the wings) at 23:50 h. This activity reached its peak at 1:50–3:30 h and ended at 3:40 h. Mated females had three oviposition periods (1.5%): the first at 23:00 h, the second from 1:30–2:00 h, and the last at 3:00 h (Fig. 1).

Fig. 1
figure 1

Observation for 12 h (18:00–6:00 h) of virgin (VF) and mated (MF) females in field cages (N = 10). First recorded activity ▲, peak activity , and last recorded activity ▼

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Attraction to Cedrela odorata Plants. Virgin females (G = 40.34; df = 1; P < 0.001), virgin males (G = 11.28; df = 1; P < 0.001), and mated females (G = 17.45; df = 1; P < 0.001) were significantly more attracted to cedar plants than to the control (dummy). It should be noted, however, that almost half of the mated females did not respond (48%, Fig. 2).

Fig. 2
figure 2

First choice response of Hypsipyla grandella in a field cage to Cedrela odorata plants. Asterisks indicate significant differences between treatments using the G test, * P < 0.05 ** P < 0.01 *** P < 0.001 significance level. N = 50. NR = indicate the number of moths that did not respond to any treatment. Cedar plant or control = dummy

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Attraction to Cedrela odorata Volatiles and Synthetic Blend. Volatiles from the headspace of plants caused significantly greater responses in virgin females (G = 18.29; df = 1; P < 0.001), virgin males (G = 10.71; df = 1; P < 0.01), and mated females (G = 6.86; df = 1; P < 0.01) than the control. However, as in the attraction bioassays of H. grandella to C. odorata plants, the number of non-responding mated females was relatively high (74%, Fig. 3).

Fig. 3
figure 3

First choice response of Hypsipyla grandella in a Y-tube olfactometer to headspace volatiles from Cedrela odorata extracts. Asterisks indicate significant differences between treatments using the G test, * P < 0.05 ** P < 0.01 *** P < 0.001 significance level. N = 50. NR = indicate the number of moths that did not respond to any treatment. Cedar extract containing volatiles from C. odorata plants or control = dichloromethane

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The synthetic blend was significantly more attractive to mated females (G = 19.53; df = 1; P < 0.001) and virgin males (G = 6.91; df = 1; P < 0.01) than the control. Virgin females, however, were not significantly more attracted to the synthetic blend than to the control (G = 1.70; df = 1; P > 0.05) (Fig. 4).

Fig. 4
figure 4

First choice response of Hypsipyla grandella in a Y-tube olfactometer to a synthetic blend that mimics the composition of Cedrela odorata headspace volatile emissions. Asterisks indicate significant differences between treatments using the G test, * P < 0.05 ** P < 0.01 *** P < 0.001 significance level. ns = non-significant difference. N = 50. NR = indicate the number of moths that did not respond to any treatment. Synthetic blend or control = dichloromethane

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Identification of Volatiles

GC–MS analysis of the volatile compounds from cedar plants revealed the presence of nine compounds. 2-Ethyl-1-hexanol and nonanal were the main components followed by α-copaene, (E)-4,8-dimethyl-1,3,7-nonatriene, α-pinene, limonene, germacrene D, β-caryophyllene, and (E)-β-ocimene (Table 1).

Discussion

In this study, we observed that H. grandella exhibited low activity during the night, despite being nocturnal insects, and that its antennae were constantly moving, which was also reported by Barradas-Juanz et al. (2016). We also investigated the attraction response of H. grandella (virgin males and mated and virgin females) to cedar volatiles. We determined the behavior of H. grandella towards the C. odorata plant volatiles to know the behavioral activities of the insects in presence of its host plant. Numerous studies have reported on interactions between insect pheromones and host plant volatiles (Landolt and Phillips 1997). Plant volatiles can signal suitable meeting sites for mating or serve to increase the response to their sex pheromone (Schmidt-Büsser et al. 2009). For example, we observed flights (of virgin as well as mated females) between 18:50 and 4:00 h, and peaked at 19:50–22:00 h. However, Gara et al. (1972) recorded flights between 0:00 and 5:00 h, and noted that no flights occurred when night temperatures dropped below 15 ºC. Furthermore, the moths can fly 31.4 km, although they won’t easily leave affected areas (Grijpma and Gara 1970; Fasoranti et al. 1982). Grijpma and Gara (1970) reported that H. grandella can be found in ground vegetation during the day, and that it walked from one tree to another. In our study, we frequently observed walking activity and recorded a peak at 21:00–22:50 h. We observed that female moths assume a calling position by exposing the abdominal glands while wing-fanning as part of their circadian rhythm of calling behavior (Barradas-Juanz et al. 2016; Levi-Zada and Byers 2021). Females remain in the position for 1.6 h to attract male conspecifics (Grijpma 1971; Samaniego and Sterringa 1976), which approximates our observed duration of 2 h. The moth’s wing-fanning behavior was first reported by Grijpma (1971). It is thought that females beat their wings to disperse their pheromones (Levi-Zada and Byers 2021). The wing-fanning behavior that we observed was similar to the behavior reported by Barradas-Juanz et al. (2016), who characterized two different types of wing-fanning – short (< 10 s) and long (> 30 s) – between 3 and 4 h. It is currently not clear whether the different types of wing-fanning also imply different functions, although both types are likely involved in pheromone dispersion (Barradas-Juanz et al. 2016).

Volatile compounds are often essential for insects to locate their hosts. To recognize their hosts, insects often use specific compounds or specific blends of ubiquitous compounds (Bruce et al. 2005; Bruce and Pickett 2011). Our results suggest that H. grandella moths are attracted by volatiles emitted by the cedar plant. Also, a synthetic blend confirmed that the identified compounds could be responsible for the attractant effect.

We demonstrated that volatile compounds from cedar plants are involved in attracting H. grandella; several studies have shown that the attack of this moth is closely related to the foliar phenology of the trees (Newton et al. 1998). Although the localization mechanism of the host plant of this species (H. grandella) is still unclear, the orientation by volatiles of the host seems to be the most likely mechanism to localize the plant (Grijpma and Gara 1970; Gara et al. 1972; Grijpma 1976; Howard 1991; Prokopy and Papaj 2001). Volatiles released by the plant can stimulate females to oviposit (Grijpma 1976), or as occurs with other lepidoptera, these could synergize response to sex pheromone (Yang et al. 2004; Xiang et al. 2019). We are thinking performance future bioassays to evaluate the effect of the sex pheromone combined with host plant volatiles.

Mated males and females of H. grandella showed a different choice response for C. odorata than virgin females (Newton et al. 1998). The fact than virgin and mated females of H. grandella respond differently to cedar volatiles may be related to many factors, for instance the insect olfactory system (Röstelien et al. 2000), or differences in the chemical composition of volatiles from young and mature foliage (Gara et al. 1972). We found that virgin females were not attracted to the synthetic blend. Possibly, the absence of germacrene D from the blend explains why the virgin female response was not significantly different for the blend. However, more studies are needed to demonstrate this suggestion. On the other hand, the high number of non-responses of the moths to the evaluated stimuli is possibly due to internal factors such as age, circadian rhythm, sexual status, egg load, or the level of hunger of the evaluated insects (Rojas 2012). In other studies, however, mated female moths responded in greater numbers than virgin female and male moths to volatile compounds (Rojas 1999), as was the case with the synthetic blend we evaluated. This may be because the insect is motivated by finding its host plant to lay its eggs.

Plants release volatile compounds such as sesquiterpenes that play an essential biological role in insect-plant interactions (Paré and Tumlinson 1999; Das et al. 2013). Sesquiterpenes (including α-copaene, β-elemene, β-caryophyllene and germacrene D) have been identified in the essential oil of terminal shoots and of mature and senescent leaves of S. macrophylla (Soares et al. 2003). Abraham et al. (2014) characterized volatile organic compounds in four different mahogany species and found various sesquiterpenes, including α-cubebene, α-copaene, β-elemene, β-caryophyllene, α-humulene, germacrene D, and δ-cadinene. The same sesquiterpenes have also been identified in various Meliaceae, such as C. odorata, Cedrela fissilis (Vell.), and Toona ciliata (M. Roem) (Maia et al. 2000). Lago et al. (2006) detected various sesquiterpenes (including α-cubebene, α-copaene, β-caryophyllene, α-humulene y germacrene D) in the essential oil of Guarea macrophylla (Vahl).

We identified nine compounds in the volatile profile of C. odorata: α-pinene, (E)-β-ocimene, 2-ethyl-1-hexanol, limonene, nonanal, (E)-4,8-dimethyl-1,3,7-nonatriene, α-copaene, β-caryophyllene, and germacrene D. A synthetic blend was prepared that mimicked the composition of C. odorata headspace volatile emissions as attractant to H. grandella moths. However, due to its unavailability, we could not include germacrene D in the synthetic blend. β-caryophyllene, one of the constituents of the synthetic blend, triggered an antennal response in female shoot-borers (Soares et al. 2003). According to Borges et al. (2022), nonanal, decanal, methyl salicylate, and β-caryophyllene elicit antennal response in male as well as female shoot-borers. The same authors found that methyl salicylate negatively affects host plant location by H. grandella. Field areas treated with this compound were less attacked by this moth, which opens the potential to integrate this compound in shoot borer pest management strategies. β-Caryophyllene and germacrene D, both identified in our volatile analyses, have been shown to strongly affect insect-plant interactions in other ecosystems (Mozuraitis et al. 2002; Köllner et al. 2008; Hare 2011; Xiao et al. 2012). Abraham et al. (2014), for instance, noted that germacrene D elicited an antenna response in female H. robusta moths. Moreover, Lago and Roque (2002) identified α-copaene, β-caryophyllene, and germacrene D in essential oil extracted from the leaves of G. macrophylla. Cedar extract also contained (E)-β-ocimene, 2-ethyl-1-hexanol and nonanal, which have been reported as antennal active compounds in female H. robusta moths (Abraham et al. 2014). Nonanal is a known oviposition attractant for the light brown apple moth Epiphyas postvittana Walker (Suckling et al. 1996), and (E)-4,8-dimethyl-1,3,7-nonatriene has been used as a lure to catch male and female Cydia pomonella (L.) (Knight et al. 2011). On the other hand, α-pinene and D-limonene have been reported to influence plant–insect interactions in the order Coleoptera (Tafoya et al. 2011; Romero-Frías et al. 2015; Sánchez-Martínez and Reséndiz-Martínez 2020).

In summary, in this work we observed that virgin and mated female shoot borer H. grandella exhibited low activity during the night, frequent antennal movement, sporadic flight activity, and short and long wing-fanning. Virgin females assumed a calling position, whereas mated females exhibited three periods of oviposition. Virgin males and females as well as mated females of H. grandella were attracted by volatiles released by the cedar plant and we identify nine volatiles in the headspace of cedar: α-pinene, (E)-β-ocimene, 2-ethyl-1-hexanol, limonene, nonanal, (E)-4,8-dimethyl-1,3,7-nonatriene, α-copaene, β-caryophyllene, and germacrene D. These compounds, except for germacrene D, were combined to prepare a synthetic blend that mimicked C. odorata volatile emissions. Male virgin and mated female H. grandella moths were attracted to the blend. Future research should be carried out to assess the potential of the synthetic blend in the field. The development of an effective attractant for the H. grandella moth would greatly improve our ability to manage this pest.