Sensilla Chaetica Research Articles

Morphology and function of ovipositorial and tarsal sensilla of female Asian citrus psyllid

AbstractThe Asian citrus psyllid (ACP), Diaphorina citri Kuwayama (Hemiptera:Psyllidae), is one of the most important pests of citrus plants worldwide, due to be the dominant vector of the devastating citrus disease Huanglongbing (HLB). Selection of suitable oviposition site is a crucial event of phytophagous insect life. Usually, the chemical and physical characteristics of a host plant were recognized by sensilla on the ovipositor and tarsi of a phytophagous insect after settling on a plant surface. In this study, the morphology, number, and distribution pattern of the ovipositorial and tarsal sensilla of adult female psyllids, were observed by scanning electron microscopy (SEM) techniques. The effect of these sensilla on oviposition was investigated by sensilla deactivation using hydrochloric acid (HCl). The results showed that sensilla trichoid (ST), sensilla chaetica (ChS) and sensilla basiconica (SB) were distributed on ovipositor, while sensilla trichoid (ST) were distributed on the external genitalia. Sensilla chaetica (ChS) and basiconica (SB) were mainly distributed on the end of the ovipositor back plate, and on the base of the sternum. On tarsus, sensilla trichoid (ST), sensilla chaetica (ChS), sensilla triangular (TeS) and sensilla ear ball (SE) were observed. However, whole tarsal segments were covered with Böhm's mane, which was arranged in a circular line. Oviposition assay revealed that a significantly lower number of eggs were laid on the Murraya paniculata seedlings confined by the psyllids with deactivated ovipositorial sensilla followed by protarsal sensilla, mesotarsal sensilla and hindtarsal sensilla. These results suggested that the ovipositorial and tarsal sensilla of citrus psyllid have an important role in the oviposition.

Morphological Effect of Gamma Radiation on the Antennal Sensory Structure of Peach Fruit Fly, Bactrocera zonata, Using Electron Microscopy

External morphology of antennal sensilla of both sexes of irradiated with sterilizing and substerilizing doses and unirradiated peach fruit fly, Bactrocera zonata were studied with scanning electron microscopy. The antenna of this fly consists of 3 segments: Basal scape, pedicel and funiculus. The antennal segments of males tended to be longer and more elongated than those of females. Eight distinct morphological types of sensilla belong to four groups. These sensilla are: trichoid sensilla type I and II, chaetica sensilla type I and II, basiconic sensilla type I, II, III and campaniform sensilla (in the pedicel of female). The changes which occurred to the antennae and their associated sensilla due to the sterilizing dose (90Gy for male and 50Gy for female) were investigated; distribution, size and position are also described.

Morphological comparison of proboscides and associated sensilla of Helicoverpa armigera and Mythimna separate (Lepidoptera: Noctuidae)

Proboscides are important feeding devices for most adult Lepidoptera and exhibit significant morphological modifications and types of sensilla associated with feeding habits. In this study the architectures of the proboscides and sensilla were compared between the cotton bollworm Helicoverpa armigera (Hübner) and the armyworm Mythimna separate (Walker) using scanning electron microscopy. The proboscides of both species consist of two elongated maxillary galeae joined by dorsal and ventral legulae, forming a food canal. The dorsal legulae in H. armigera disappear a short distance from the proboscis apex, whereas those in M. separate exist up to the apex. Three types of sensilla are present on the proboscides of both species: sensilla chaetica, basiconica, and styloconica. The morphological differences of the sensilla mainly concern the sensilla styloconica, whose styli have six to seven smooth-edged ridges in H. armigera but six serrate-edged ridges in M. separate. No significant sexual dimorphism was found in the proboscides and sensilla of both species except for the length of the zone without the dorsal legulae in H. armigera. The morphological similarities and differences of the proboscides and sensilla between the two species are briefly discussed.

Expression of odorant-binding proteins in mouthpart palps of the desert locust Schistocerca gregaria.

Odorant‐binding proteins (OBPs) are essential molecular elements of the insect chemosensory system, which is composed of the antennae and the mouthpart palps (maxillary and labial). In this study, we have analysed the expression and the sensilla specificity of 14 OBP subtypes in the palps of the desert locust Schistocerca gregaria. The locust palps comprise only a low number of sensilla basiconica but a high number of sensilla chaetica. Employing a variety of approaches, we found that only a subset of the antennal OBP repertoire was expressed in both palp types. These OBPs were previously shown to be expressed either in sensilla basiconica or sensilla chaetica of the antennae. Comparing the expression pattern in the two chemosensory organs revealed similarities and differences; most remarkably, two OBP subtypes, OBP6 and OBP8, were found in both sensilla types on palps, whereas on the antennae they were solely expressed in one sensillum type. Together, the data indicate a differential, but partly overlapping, expression of OBPs in the two sensilla types of the palps. The differences in the expression pattern of OBP subtypes between antennae and palps might be indicative for distinct functions of the OBPs in the two chemosensory organs.

The structure and morphologic changes of antennae of Cyrtorhinus lividipennis (Hemiptera: Miridae: Orthotylinae) in different instars.

Cyrtorhinus lividipennis Reuter (Hemiptera: Miridae: Orthotylinae), including nymphs and adults, are one of the dominant predators and have a significant role in the biological control of leafhoppers and planthoppers in irrigated rice. In this study, we investigated the antennal morphology, structure and sensilla distribution of C. lividipennis in different instars using scanning electron microscopy. The antennae of both five different nymphal stages and adults were filiform in shape, which consisted of the scape, pedicel and flagellum with two flagellomeres. There were significant differences found in the types of antennal sensilla between nymphs and adults. The multiporous placodea sensilla (MPLA), basiconica sensilla II (BAS II), and sensory pits (SP) only occurred on the antennae of adult C. lividipennis of both sexes. Moreover, there was chaetica sensilla III (CHA III) only observed in males. Sixteen types of antennal sensilla were recorded altogether. They were microtrichia sensilla (MIC), three types of trichoidea sensilla (TRI I-III), three types of chaetica sensilla (CHA I-III), three types of basiconica sensilla (BAS I-III), two types of coeloconica sensilla (COE I and COE II), placodea sensilla (PLA), campaniform sensilla (CAM), MPLA, and SP. In the five different nymphal stages of C. lividipennis, the length of their antennae was significantly increased with the increase of the instar, as well as the number of the TRI II and TRI III. Moreover, sexual dimorphism usually occurred not only in the distribution (CHA III and SP) and the number of antennal sensilla (MIC, BAS II, TRI II, TRI III and MPLA), but also in the length of flagellum (F1 and F2). The possible functions of antennal sensilla are discussed. Those observations could contribute to a better understanding of the development of the olfactory system, and facilitate future studies on the antennal functions in C. lividipennis.

Antennal Morphology and Localization of a Pheromone-Binding Protein of Lobesia botrana (Denis & Schiffermüller) (Lepidoptera: Tortricidae).

In the sensory system of insects, olfactory sensilla constitute important functional elements for discriminating odors. Therefore, we used light microscopy and scanning electron microscopy to investigate the morphology and distribution of sensilla in the antennae of Lobesia botrana (Denis & Schiffermüller). In addition, we studied the expression of the gene encoding for pheromone-binding protein 1 (LbotPBP1) by in situ hybridization. Lobesia botrana antennae are filiform and are subdivided into three segments: scape, pedicel, and flagellum. The number of flagellum and their overall length were significantly higher and longer in males than in females. Six morphological types of sensilla (trichodea, chaetica, coeloconica, auricillica, basiconica, and styloconica) were identified on the antennae of both sexes. Trichodea sensilla were the most abundant on the antennae of L. botrana, and three subtypes, discerned by their lengths, were observed. However, sensilla trichodea subtype III was only present in male antennae. Moreover, LbotPBP1 expression was restricted to this type of sensilla, thus confirming its olfactory role, specifically under the context of sexual pheromone perception.

Sensilla on six olfactory organs of male Eogystia hippophaecolus (Lepidoptera: Cossidae).

Eogystia hippophaecolus (Hua et al.) is an important boring pest that primarily damages sea buckthorn, causing large ecological and economic damages in China. In this study, we used scanning electron microscopy to investigate the sensilla on six olfactory tissues of male E. hippophaecolus: antennae, labial palps, external genitals, propodeum, mesopodium, and metapedes. On the antennae, two types of sensillum trichodea, two types of sensillum basiconica, a type of sensillum coeloconica, and Böhm's bristles were found. The labial palps had sensilla trichodea and chaetica. On the external genitals, three types of sensilla trichodea were the only sensilla. Böhm bristles were found on the base of the tibia and at the terminus of the tarsus. Most sensilla were distributed on the tarsus of the three pairs of legs, and notably, a majority of which were sensilla trichodea and sensilla chaetica, were on the pretarsus and telotarsus of the three pairs of legs. In this study, the distribution, number, type, and morphology of the sensilla on six olfactory organs of E. hippophaecolus were determined, which established the foundation for a future immunohistochemical search of olfactory proteins. RESEARCH HIGHLIGHTS: Eogystia hippophaecolus is a serious pest of seabuckthorn. The distribution, number, type, and morphology of olfactory sensilla of E. hippophaecolus are determined. The function and distribution of sensilla are compared with those of other insects.

Antennal morphology and ultrastructure of male and female Ptinus japonicus Reitter, 1877 (Coleoptera: Ptinidae)

Ptinus japonicus Reitter is a subversive pest of stored-product throughout the world. Scanning electron microscopy (SEM) was used to observe the antennal morphology and sensilla ultrastructure in both sexes of this pest. The antennae of P. japonicus are filiform that comprised scape, pedicel and flagellum with nine flagellomeres. Six types of sensilla were identified, including Böhm sensilla (Bs), two subtypes of sensilla basiconica (SbI-II), sensilla styloconica (Sst), four subtypes of sensilla chaetica (SchI-IV), sensilla coeloconica (Sco), two subtypes of sensilla trichodea (St I-II). The antennae of Ptinus japonicus exhibit obvious sexual dimorphism in antennal size, the type and size of the sensilla, the distribution and abundance of the sensilla. The length of antennae in males was significantly longer than these in females. Sensilla trichodea I (StI) were found only in males. The number of sensilla basiconica (Sb) and Sensilla chaetica IV (SchIV) in females were higher than in males. Bs in females was longer than in males. The results of this study provide theoretical basis and material for the further study on mating and habitant searching, and may provide a new sight into semiochemical-based control of spider beetles.

Ultrastructure of Female Antennal Sensilla of an Endoparasitoid Wasp, Quadrastichus mendeli Kim & La Salle (Hymenoptera: Eulophidae: Tetrastichinae).

The antennal sensilla of female Quadrastichus mendeli Kim & La Salle (Hymenoptera: Eulophidae: Tetrastichinae) were observed with scanning and transmission electron microscopy in this study. The antenna of Q. mendeli was geniculate, and the flagellum was composed of seven subsegments. Six distinct types of sensory receptors were observed, including sensilla basiconic capitate peg, sensilla böhm, sensilla chaetica, sensilla campaniformia, sensilla placodea and sensilla trichodea. Sensilla basiconic capitate pegs were found on the flagellomeres, and Böhm sensilla were found on the basal part of scape and the pedicel. Two morphological subtypes of sensilla chaetica were found on the antennae, and sensilla campaniformia were only found on the pedicel. Sensilla placodea were divided into two morphological subtypes that were found on the flagellomeres. Sensilla trichodea were found on the 2nd-6th flagellomere. By comparison to existing antennal sensilla, it was found that sensilla basiconic capitate peg, sensilla chaetica, sensilla placodea and sensilla trichodea were the most common sensilla of the parasitoids of Eulophidae. The external and internal morphology, types, number, distribution, length, and width of these sensilla were described, and their possible functions are discussed in conjunction with the host-detection behavior. Future studies on the host location mechanisms in Q. mendeli will be facilitated by these observations.

Single Sensillum Recordings for Locust Palp Sensilla Basiconica.

The palps of locust mouthparts are considered to be conventional gustatory organs that play an important role in a locust's food selection, especially for the detection of non-volatile chemical cues through sensilla chaetica (previously named terminal sensilla or crested sensilla). There is now increasing evidence that these palps also have an olfactory function. An odorant receptor (LmigOR2) and an odorant-binding protein (LmigOBP1) have been localized in the neurons and accessory cells, respectively, in the sensilla basiconica of the palps. Single sensillum recording (SSR) is used for recording the responses of odorant receptor neurons, which is an effective method for screening active ligands on specific odorant receptors. SSR is used in functional studies of odorant receptors in palp sensilla. The structure of the sensilla basiconica located on the dome of the palps differs somewhat from the structure of those on the antennae. Therefore, when performing an SSR elicited by odorants, some specific advice may be helpful for obtaining optimum results. In this paper, a detailed and highly effective protocol for an SSR from insect palp sensilla basiconica is introduced.

Ultrastructure of the sensilla on antennae and mouthparts of larval and adult Plutella xylostella (Lepidoptera: Plutellidae)

Plutella xylostella (L.) (Plutellidae) is an important agricultural pest throughout the world. In this study, the morphology of antennal and mouthpart sensilla in the larvae and adults of P. xylostella (L.) was observed by using a scanning electron microscope. The larval antennae possess six sensilla basiconica, two sensilla chaetica and one sensillum styloconicum. Larval mouthparts possess six types of sensilla: sensilla chaetica, sensilla digitiformia, sensilla epipharyngeal, sensilla basiconica, sensilla styloconica and sensilla placodea. In the adult, seven types of sensilla are found on the antennae in males and six types of sensilla (sensilla basiconica absent) occur in females. Sexual dimorphism is also found in the number and size of these sensilla on the antennae of adults. We describe for the first time the five types of sensilla on the mouthparts of the adult of P. xylostella. This study provides useful information for further research into the function of these sensilla, and better understanding the behavioral mechanisms involved in pest control.

The Odorant Binding Protein 6 Expressed in Sensilla Chaetica Displays Preferential Binding Affinity to Host Plants Volatiles in Ectropis obliqua.

The monophagous tea geometrid Ectropis obliqua selectively feed on tea plants, requiring the specialized chemosensory system to forage for certain host. A deep insight into the molecular basis would accelerate the design of insect-behavior-modifying stimuli. In the present study, we focused on the odorant-binding protein 6 (EoblOBP6) with the high abundance in legs transcriptome of E. obliqua moths. qRT-PCR coupled with western blot analyses revealed the dual expression pattern of EoblOBP6 in antennae and legs. Cellular immunolocalization indicated that EoblOBP6 was predominantly labeled in the outer sensillum lymph of uniporous sensilla chaetica, which is not innervated by sensory neurons. No specific staining was observed in other sensillum types. The fluorescence competition assay showed a relatively narrow binding spectrum of recombinant EoblOBP6. EoblOBP6 could not only bind with intact tea plant volatiles benzaldehyde but also display high binding ability to nerolidol and α-farnesene which are tea plant volatiles dramatically induced by herbivore infestation. Besides, EoblOBP6 tightly bound to the aversive bitter alkaloid berberine. Taken together, EoblOBP6 displayed an unusual expression in sensilla chaetica, exhibited the potential involvement in olfaction and gustation, and may play a functional role in host location of female E. obliqua moths.

Odorant Binding Proteins of the Desert Locust Schistocerca gregaria (Orthoptera, Acrididae): Topographic Expression Patterns in the Antennae.

Odorant binding proteins (OBPs) enriched in the sensillum lymph are instrumental in facilitating the transfer of odorous molecules to the responsive receptors. In Orthopteran locust species, an in-depth understanding of this important soluble protein family is still elusive. In a previous study, we have demonstrated that the repertoire of locust OBPs can be divided into four major clades (I–IV) on the phylogenetic scale and for representatives of subfamily I-A and II-A a distinct sensilla-specific expression pattern was determined. In this study, by focusing on a representative locust species, the desert locust Schistocerca gregaria, we have explored the antennal topographic expression for representative OBPs of other subfamilies. First, subtypes of subfamily III-A and III-B were exclusively found in sensilla chaetica. Then, a similar expression pattern in this sensillum type was observed for subfamily I-B subtypes, but with a distinct OBP that was expressed in sensilla coeloconica additionally. Moreover, the atypical OBP subtype from subfamily IV-A was expressed in a subpopulation of sensilla coeloconica. Last, the plus-C type-B OBP subtype from subfamily IV-B seems to be associated with all four antennal sensillum types. These results profile diversified sensilla-specific expression patterns of the desert locust OBPs from different subfamilies and complex co-localization phenotypes of distinct OBP subtypes in defined sensilla, which provide informative clues concerning their possible functional mode as well as a potential interplay among OBP partners within a sensillum.

Morphological characterization and distribution of antennal sensilla of irradiated female mosquito, Culex pipiens (Diptera: Culicidae) with gamma radiation

Culex pipiens mosquito is considered as one of the most dangerous insect pests that affect humans and animals worldwide as they transmit epidemic and fatal diseases. Antennal structure of the normal and irradiated female mosquito, Culex pipiens was examined using scanning electron microscopy to study the changes in the antennal segments and associated sensilla when 1–2 days old pupae were irradiated at low dose (20Gy), moderate (40Gy) and high (80Gy). The antenna of the female C. pipiens is filamentous in shape and consisted of scape (S), pedicel (P) and flagellum (F) which has 13 flagellomeres. Various types of sensilla with distinct morphological characters and morphometric features were recorded. Sensilla trichodea are the most abundant sensilla observed, with other types like sensilla chaetica, sensilla basiconica (grooved peg), sensilla coeloconica and sensilla ampullaceal. Morphological changes were recorded to the antennal segments and their associated sensilla due to the doses of gamma radiation (20, 40 and 80Gy) such as shrinkage, curved of sensilla into different directions, irregularity in shape, swollen in some parts and some sensilla gathered into dense collections. These changes were obviously demonstrated at 80Gy and 40Gy doses whereas little malformations were observed when the pupae irradiated with 20Gy dose. The antenna has lost most of different types of sensilla and some of its segments became transparent with high and moderate doses. These changes also recorded in the distal 13th flagellomeres segments with the three used doses of gamma radiation compared to control ones.

Ultrastructure of sensilla on the maxillary and labial palps of the adult Xylotrechus grayii (Coleoptera: Cerambycidae).

The longhorn beetle, Xylotrechus grayii (White, 1855), is a serious woodborer pest of many economic and medicinal plants. The maxillary and labial palps of X. grayii adults are strongly involved in mating and host selection behaviors. To further develop control strategies of the pest, sensilla on the palps were studied with scanning and transmission electron microscopy. Five types of sensilla (nine subtypes) were identified: sensilla styloconica (four subtypes, Sty1-4), sensilla chaetica (2 subtypes, SC1-2), sensilla digitiformia (Di), sensilla basiconica (Ba), and Böhm sensilla (Bm). Sty and Ba have dendrites and wall pores. SC2 has a tubular body. Di exist only on the terminal segment of the maxillary palp. Sty numbers are the highest in both palps, followed by SC. The number of sensilla in the maxillary palp is significantly higher than that in the labial palp. There are no significant sexual differences in sensilla types, characteristics, distribution, and quantity in the maxillary or labial palps, except for the number of SC sensilla on the maxillary palp. Sensilla functions were compared with correlative sensilla in previous studies. Sty may be sense gustation and olfaction stimulation, while SC are mechanoreceptors. Ba are likely thermo- and hygroreceptors. The phylliform depression area formed by Di may exert multiple functions. Bm may sense palp position and movement. These results contribute to the study of the electrophysiology and behavior mechanisms of X. grayii.

Distribution and Ultrastructure of Sensilla onUlomoides dermestoides(Coleoptera: Tenebrionidae) Antennae1

Abstract Ulomoides dermestoides (Chevrolat) (Coleoptera, Tenebrionidae) is a storage pest and an insect of medical importance. Since this insect lives in the dark and does not emit sounds, its sensation of the external environment is based primarily on sensilla, so it could serve as an ideal model for the study of these structures. Scanning electron microscopy was used to identify the types of antennal sensilla of male and female U. dermestoides and to characterize the function of different types of sensilla observed. The bead-like antenna of U. dermestoides is composed of a scape (one segment), a pedicel (one segment), and a flagellum (nine segments). Six types of sensilla were observed on the antennae of both females and males: the sensilla trichoidea (subtypes I, II, and III), sensilla chaetica (subtypes I, II, and III), sensilla basiconica, basiconic mastoid sensillum, sensilla styloconica (subtypes I and II), and the Bohm bristle. The Student's t test was used to compare the distribution of these sen...

Putative adhesive setae on the walking legs of the Paleotropical harvestman Metibalonius sp. (Arachnida: Opiliones: Podoctidae)

We provide a first scanning electron microscopy examination of the Paleotropical harvestman family Podoctidae (Opiliones: Laniatores), focusing on the distitarsus of the legs of Metibalonius sp. Distitarsi I and II are mostly equipped with olfactory sensilla chaetica with wall pores, while those of legs III and IV have gustatory sensilla chaetica with a tip pore, ventral trichomes with ovate tips (non-sensory) and a type of spatulate seta. Spatulate setae are present in adults of both sexes, with no apparent sexual dimorphism, but they are absent in the nymph. Seven of these setae are inserted on the frontal surface of the last tarsomere of legs III and IV, with the tips oriented ventrally. Each seta has an s-shaped socketed shaft, which terminates distally in a spatula-shaped structure. The distribution of spatulate setae, restricted to legs III and IV (walking legs), the position on the distitarsi, and the typical spatulate shape suggest an adhesive function for these structures. Morphology and position suggest that the socketed spatulate setae of Metibalonius sp. and the previously reported scopular spatulate setae of other harvestmen constitute two distinct types of adhesive structures, highlighting the diversity of adhesive structures in Laniatores. Future investigations about the natural history of this species and internal morphology of spatulate setae are necessary to test further functional hypotheses and to determine their behavioral role.

Morphology and distribution of antennal, maxillary palp and labial palp sensilla of the adult bruchid beetles,Callosobruchus chinensis(L.) (Coleoptera: Bruchidae)

AbstractCallosobruchus chinensis(Coleoptera: Bruchidae) is the main pest of mungbean (Vigna radiata) especially during seed storage. To provide background information for chemical ecology studies, we examined the externalmorphology of the antennal, maxillary palp and labial palp sensilla of this insect with scanning electron microscopy. The antennae of adultC. chinensisexhibited sexual dimorphism. The female have serrated antenna and the male have pectinate morphology. Eight sensillum types have been observed on the antennae of both sexes: Böhm bristles, sensilla chaetica, sensilla trichodea, sensilla basiconica, sensilla styloconicum, sensilla coeloconica, sensillum campaniformia, sensilla cavity. There are no differences on the morphology of maxillary palp and labial palp between female and male. Four different types of sensilla are present on the labial and maxillary palps of both sexes ofC. chinensis: sensilla chaetica, sensilla twig basiconica, sensilla basiconica, sensilla digitiformia.

Distribution and ultrastructure of the antennal sensilla of the grape weevil Naupactus xanthographus (Coleoptera: Curculionidae).

The grape weevil, Naupactus xanthographus Germar (Coleoptera: Curculionidae), is a polyphagous insect native to southern South America that causes considerable damage in grape and other fruit species. In this study, the morphology and ultrastructure of the antennae and the antennal sensilla of N. xanthographus were investigated using scanning electron microscopy and transmission electron microscopy. The antennae consist of a scape, a pedicel, a funicle, and a zone called the "club," which are all formed by a total of 12 antennomers. Different types of sensilla were observed: sensilla trichoidea, sensilla chaetica type 1 and 2, sensilla basiconica, and sensilla rod-like. No sexual dimorphism was observed. The possible functions of the sensilla are discussed in relation to their morphology and ultrastructure.

Antennal sensilla of Didesmococcus koreanus Borchsenius (Hemiptera: Coccoidea: Coccidae) in different instars

To better understand the morphology, structure and function of antennae and antennal sensilla of Didesmococcus koreanus Borchsenius, family Coccidae (Hemiptera: Coccoidea), in the first-instar nymph, adult female and adult male stages, we investigated these stages using light microscopy and scanning and transmission electron microscopy. The results show that the antennae of first-instar nymphs and adult females are similar in morphology and sensilla distribution. The antennae of first-instar nymphs comprise six segments, and each antenna has 24 sensilla of six types: one campaniform sensillum (Ca), eleven smooth aporous trichoid sensilla (St), two basiconic sensilla (Ba), five multiporous pegs (Mp), two smooth multiporous pegs (sMp) and three chaetica sensilla (Ch). The antennae of adult females have six or seven segments, and each antenna has 25 sensilla of the same six types as the first nymphs, but with twelve sensillum St. The antennae of adult males are more developed, and each antenna contains 155–213 sensilla of eight types: 16–34 St, 1 Ca, 2–5 Ba, 1 Johnston’s organ (Jo), 3 knob sensilla (Ks), 5 Mp, 2 sMp and 120–181 multiporous sensilla trichodea (Mt). Based on our analysis, we discuss the possible functions of the antennae and antennal sensilla of D. koreanus in different instars.