The antennae of neopseustid moths: Morphology and phylogenetic implications, with special reference to the sensilla (Insecta, Lepidoptera, Neopseustidae)

The antennae of Lepidoptera Neopseustidae were examined with the scanning electron microscope. The studied species, Nematocentropus cfr. omeiensis, Neopseustis meyricki, Synempora andesae, Apoplania valdiviana and Apoplania penai possess nine types of antennal flagellum sensilla: multiporous large sensilla basiconica, multiporous thin sensilla basiconica, multiporous small sensilla basiconica, multiporous sensilla trichodea, multiporous sensilla coeloconica; uniporous sensilla chaetica; aporous sensilla chaetica, aporous stylus-shaped sensilla chaetica, aporous sensilla styloconica.The multiporous sensillum type here termed “multiporous large sensillum basiconicum” is unknown from other Lepidoptera and probably constitutes an autapomorphy of the family Neopseustidae. This sensillum type is remarkable by having a single base in female Apoplania and Synempora while in male Apoplania it has a bifid or trifid base, and in male Synempora it is composed of two or three incompletely separated hairs. This may be the first recorded example of a sexually dimorphic lepidopteran sensillum type. The stylus-shaped sensillum chaeticum is a primitive type which occurs only in some lower Lepidoptera.     

Sensilla on the larval antennae and mouthparts of the european sunflower moth,Homoeosoma nebulella Den. and Schiff. (Lepidoptera : Pyralidae)

Sensilla on the larval antennae and mouthparts of Homoeosoma nebulella (Lepidoptera : Pyralidae) were examined with a scanning electron microscope. The antenna possesses 2 aporous sensilla chaetica, one aporous sensillum styloconicum, 3 large multiporous sensilla basiconica, and 3 small multiporous s. basiconica. The labrum bears 12 aporous s. chaetica and the epipharynx has 6 aporous flattened s. chaetica and 4 epipharyngeal sensilla. It is the first time that more than 2 epipharyngeal sensilla have been reported in caterpillars. The mandible possesses 4 aporous s. chaetica. Each galea bears 2 uniporous s. styloconica, 2 small aporous s. basiconica, one s. campaniformium and 3 aporous s. chaetica. Each maxillary palp bears a distal group of 8 uniporous s. basiconica, one large s. digitiformium and one pore plate. Minute grooves exist on the s. digitiformium. Each labial palp possesses one aporous s. chaeticum and one aporous s. styloconicum. An attempt has been made to identify the function of sensilla by comparing them with those of other larvae.     

Morphology and untrastructure of the antennal sense organs in tenebrionid larvae <Emphasis Type="Italic">Tenebrio molitor</Emphasis> L. and <Emphasis Type="Italic">Zophobas rugipes</Emphasis> Kirsch (Coleoptera: Tenebrionidae)

The distribution, external morphology, and ultrastructure of various types of sensilla in the antennae of tenebrionid larvae Tenebrio molitor and Zophobas rugipes are studied by means of scanning and transmission electron microscopy. On the antennae of T. molitor there are sensilla of four basic morphological types: basiconic, styloconic, trichoid, and papillate sensilla. On the antennae of Z. rugipes, in addition to the aforementioned ones, there are placoid sensilla. Ultrastructure points to olfactory function of basiconic and placoid sensilla. Other sensillum types are contact chemoreceptors.     

Morphology and distribution of sensilia on the cephalic appendages,tarsi and ovipositor of the European sunflower moth,Homoeosoma nebulella Den. & Schiff. (Lepidoptera : Pyralidae)

Sensilla on male and female antennae, mouthparts, tarsi and on the ovipositor of the European sunflower moth, Homoeosoma nebulella (Lepidoptera : Pyralidae) have been investigated by means of scanning electron microscope. The antennal flagellum bears multiporous sensilla (trichodea, basiconica, auricillica, coeloconica) and uniporous sensilla (chaetica, styloconica). A sexual dimorphism is found in the numbers of sensilla trichodea (8300 in males and 6000 in females) and sensilla basiconica (1150 in females and 650 in males). The proboscis has uniporous s. styloconica, multiporous s. basiconica (described for the first time in Lepidoptera), aporous s. chaetica, and some rare uniporous s. basiconica in the food channel. The labial palp-pit organ houses multiporous s. coeloconica, while the maxillary palps have aporous sensilla. Tarsi of both sexes possess one type of aporous s. chaetica and 2 types of uniporous s. chaetica. There is no significant difference in the distribution of tarsal sensilla between males and females. The surface of each lobe of the ovipositor is covered by 3 types of aporous s. trichodea of different lengths and about 60 multiporous s. basiconica, sensilla unusual on the ovipositor of Lepidoptera. The role of sensilla in the oviposition site selection is discussed.     

Morphology and ultrastructure of the chemosensory sensilla of larval antennae in the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae)

The allocation, external morphology, and ultrastructure of various types of sensilla on the antennae have been studied in the larvae of the Colorado potato beetle (Leptinotarsa decemlineata Say) using scanning and transmission electron microscopy. It has been found that the larval antenna bears 11 sensilla of four morphological types: trichoid (five), basiconic (four), styloconic (one), and antennal cone (one). It has been shown that the sensilla are innervated by 2?C7 receptor cells: trichoid, by 2?C4 receptor cells; basiconic, by 4?C7 receptor cells; and styloconic, by 6 receptor cells. Judging by the specifics of their ultrastructural organization, the function of these sensilla is gustatory. The antennal cone is innervated by two receptor cells, and its function is olfactory.     

Functional anatomy of sensory structures on the antennae of Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae)

The ultrastructure and distribution of sensilla on the antennae of the cabbage stem flea beetle, Psylliodes chrysocephala, were investigated using scanning and transmission electron microscopy techniques. Eight different sensillar types were distinguished. These were; hair plate sensilla, sensilla chaetica, three types of sensilla trichodea, sensilla basiconica, grooved peg sensilla and styloconic sensilla. The sensilla chaetica are known to be gustatory receptors. Ultrastructure indicates that the hair plate sensilla and sensilla trichodea type one are probably mechanoreceptors, whilst the sensilla styloconica are probably thermo-hygro receptors. These thermo-hygroreceptors are unusual in that they are innervated by two sensory cells (one hygroreceptor and one thermoreceptor) rather than the more usual triad. The remaining four sensillar types all have a porous hair shaft, indicating an olfactory role. One of these (the grooved peg sensillum) may also have a thermoreceptive function. No sexual dimorphism was found in the structure, number or distribution of the antennal sensilla.     

Antennal sensilla of male and female of the nut weevil,Curculio nucum Linnaeus, 1758 (Coleoptera: Curculionidae)

The nut weevil, Curculio nucum (Linnaeus, 1758), is the main pest in hazelnut orchards (Corylus avellana L.). Semiochemicals are interesting bio control tools that could be used to manipulate the pest behaviour and to control pest populations. The study of the sensorial equipment of the insect antennae provides information on the importance of olfaction in the adult life for host plants and mate findings as well as on the putative other senses. Before electrophysiological investigation, the knowledge of antennae equipment is also necessary. The aim of this study is to determine the types, number and location of sensilla on the antennae of male and female adult C. nucum in order to determine their implication in seeking a sexual partner and a host plant. The 12-segmented antenna comprises a scape, a 7-segmented funicle and a 4-segmented club. Out of the nine sensillum types listed, three are present on the scape and the funicle and seven types on the club which gathers 71–73% of the total of sensilla. Tactile aporous sensilla chaetica C1, gustatory uniporous sensilla chaetica C2, olfactory multiporous sensilla basiconica B1 and B2 are found on both the dorsal and ventral surfaces of the club in both sexes. Thermo-hygroreceptive dome-shaped sensilla D, olfactory multiporous sensilla basiconica B3 and olfactory multiporous fluted sensilla basiconica F are found exclusively on the ventral surface of the club, suggesting that these sensilla are utilized in host plant acceptance during antennal tapping. The sexual dimorphism concerns only the numbers of sensilla chaetica C1 and sensilla basiconica B2.     

Scanning Electron Microscopy Study of the Antennal Sensilla of <Emphasis Type="Italic">Monema flavescens</Emphasis> Walker (Lepidoptera: Limacodidae)

Monema flavescens Walker (Lepidoptera: Limacodidae) is a serious polyphagous defoliator. Using scanning electron microscopy, the external morphology of the antennal sensilla of this pest was examined for a better understanding of the mechanisms of insect-insect and insect-plant chemical communications. The antennae of M. flavescens were filiform in shape, and 11 morphological types of sensilla were found in both sexes. Six types of likely chemosensory sensilla were identified: uniporous sensilla chaetica, multiporous sensilla trichodea, and four types of multiporous sensilla basiconica. The sensilla identified as likely mechanoreceptors included two subtypes of aporous sensilla chaetica, aporous sensilla coeloconica, aporous sensilla styloconica, and Böhm’s bristles, whereas the position of the antennae was monitored by Böhm’s bristles.     

Morphological and ultrastructural characterization of olfactory sensilla in Drosophila suzukii: Scanning and transmission electron microscopy

Drosophila suzukii is a serious horticultural and quarantine pest, damaging various berry crops. Although the active use of olfactory communication in D. suzukii is well-known, their olfactory sensory system has not been comprehensively reported. Therefore, the present study was carried out to understand the morphology, distribution and ultrastructure of olfactory sensilla present in the antennae and maxillary palps of D. suzukii, through scanning electron microscopy (SEM) and transmission electron microscopy (TEM). The olfactory sensilla on the antennae of D. suzukii in both sexes could be classified into three major morphological types, basiconic, trichoid and coeloconic sensilla, according to their shapes. The antennal basiconic sensilla were further divided into three subtypes and the antennal trichoid sensilla into two subtypes, respectively, according to the size of individual sensillum. In contrast to the antennal olfactory sensilla showing diverse morphology, basiconic sensilla was the only type of olfactory sensilla in the maxillary palps of D. suzukii. The basiconic sensilla in the maxillary palps could be further classified into three subtypes, based on their size. Our SEM and TEM observations indicated that multiple nanoscale pores are present on the surface of all types of olfactory sensilla in the antennae and maxillary palps, except coeloconic sensilla. The difference in the morphological types and the distribution of olfactory sensilla suggests that their olfactory functions are different between antennae and maxillary palps in D. suzukii. The results of this study provide useful information for further studies to determine the function of olfactory sensilla in D. suzukii and to understand their chemical communication system.     

Morphological and functional heterogeneity in olfactory perception between antennae and maxillary palps in the pumpkin fruit fly,Bactrocera depressa

The morphology and ultrastructure of the olfactory sensilla on the antennae and maxillary palps were investigated through scanning electron microscopy (SEM) and transmission electron microscopy (TEM), and their responses to five volatile compounds were measured using electroantenogram (EAG) and electropalpogram (EPG) techniques in the pumpkin fruit fly, Bactrocera depressa (Shiraki; Diptera: Tephritidae). Male and female B. depressa displayed distinct morphological types of olfactory sensilla in the antennae and maxillary palps, with predominant populations of trichoid, basiconic, and coeloconic sensilla. Basiconic sensilla, the most abundant type of olfactory sensilla in the antennae, could be further classified into two different types. In contrast, the maxillary palps exhibited predominant populations of a single type of curved basiconic sensilla. High‐resolution SEM observation revealed the presence of multiple nanoscale wall‐pores on the cuticular surface of trichoid and basiconic sensilla, indicating that their primary function is olfactory. In contrast, coeloconic sensilla displayed several longitudinal grooves around the sensillum peg. The TEM observation of individual antennal olfactory sensilla indicates that the basiconic sensilla are thin‐walled, while the trichoid sensilla are thick‐walled. The profile of EAG responses of male B. depressa was different from their EPG response profile, indicating that the olfactory function of maxillary palps is different from that of antennae in this species. The structural and functional variation in the olfactory sensilla between antennae and maxillary palps suggests that each plays an independent role in the perception of olfactory signals in B. depressa.     

灰茶尺蠖成虫触角及幼虫头部感器超微结构

【目的】明确灰茶尺蠖Ectropis grisescens成虫触角及幼虫头部感器的种类、形态、数量和分布,以探讨灰茶尺蠖的行为机制。【方法】利用扫描电镜技术观察灰茶尺蠖雌、雄成虫触角和5龄幼虫头部感器的超微结构。【结果】灰茶尺蠖成虫触角上分布有8种感器,分别是栓锥形感器、耳形感器、毛形感器(Ⅰ-Ⅳ)、B?hm氏鬃毛、腔锥形感器(Ⅰ和Ⅱ)、鳞形感器、锥形感器(Ⅰ和Ⅱ)和刺形感器。其中,栓锥形感器仅分布在雌蛾触角上,耳形感器、毛形感器(STⅠ-Ⅲ)仅分布在雄虫触角上。5龄幼虫触角上着生1个栓锥形感器、1个锥形感器和2个刺形感器;上唇着生有6对刺形感器,内唇着生有3对刺形感器和1对指形感器;上颚基部外侧着生有2个刺形感器;下颚及下颚须着生有5个刺形感器、9个锥形感器和2个栓锥形感器;下唇须着生有1个锥形感器和1个刺形感器;吐丝器前端着生有1对刺形感器。【结论】灰茶尺蠖雌、雄成虫触角感器存在性二型性,且雄虫上感器种类和数量较多,据此推测雄虫感受寄主植物或性信息素的能力较强;幼虫头部感器具有嗅觉和味觉功能,在其判断食物的种类和适应性等生态行为中发挥重要作用。     

Sensilla on the mouthparts and antennae of the elephant louse,Haematomyzus elephantis piaget (Phthiraptera: Haematomyzidae)

The labial palpus of the elephant louse Haematomyzus elephantis has six sensilla that represent three different types: trichoid, basiconic, and styloconic. Two rows of basiconic sensilla are situated on the dorsal and ventral surfaces of the rostrum, and each row consists of three sensilla. Male and female antennae have 15–17 trichoid sensilla situated on the scape, pedicel, and three antennal annuli. Both sexes have two sensilla basiconica on the dorsal surface of the pedicel near the junction of the scape and pedicel. Two coeloconic (tuft) sensilla are situated on the antennae of both sexes, one sensillum on each of the last two annuli. There are three plate organs, two on the last annulus and one on the penultimate annulus of the male and female antennae. Sexual dimorphism is exhibited in the male and female antennae, in that the male has about twice as many sensilla basiconica on the apex of the last annulus as does the female. The total number of sensilla basiconica on the apex of the male antennae is at least two times the number that is known to be present in any other species of lice. © 1992 Wiley-Liss, Inc.     

Mate recognition and antennal morphology of Octodonta nipae (Coleoptera: Chrysomelidae) adults

The nipa palm hispid beetle, Octodonta nipae (Maulik) has been killing palm trees since its introduction into Hainan province, China, from Malaysia in 2001. It continues to spread within Hainan province, northeast to Fujian province, and northwest to Yunnan province within China. Knowledge on signals involved in mate location and recognition could help develop effective integrated pest management programs. In the present study, we first experimentally proved that antennae were essential in success of O. nipae mating. We then excised various segments/flagellomeres of adult male and female antennae and observed their mating behavior. Results revealed that the 5th to 9th flagellomeres, especially those of males, were important for the mating success. Finally, in an attempt to elucidate the types of antennal sensilla accountable for the mating success, morphology of O. nipae antennae was studied in detail with scanning electron microscopy. Six types of sensilla were distinguished: aporous sensilla trichodea (T1), multiporous sensilla trichodea (T2), aporous sensilla chaetica (Ch1), uniporous sensilla chaetica (Ch2), multiporous sensilla basiconica (B), and Böhm sensilla (Bm). Aporous sensilla trichodea is the most abundant; multiporous sensilla trichodea and sensilla basiconica are considered as olfactory receptors, and uniporous sensilla chaetica as gustatory receptor. Importance of flagellomeres 5–9 in mating success seemed to correspond to the abundance of sensilla on these segments.     

木瓜榕传粉榕小蜂雌蜂触角感器的分布和超微形态

为探索木瓜榕传粉榕小蜂Ceratosolen emarginatus寄主定位机制, 应用扫描电镜和透射电镜观察了其雌蜂触角感器的类型、 分布和超微形态。结果显示： 木瓜榕传粉榕小蜂雌蜂触角呈膝状, 由柄节、 梗节和11个鞭小节组成的鞭节组成, 第3鞭小节着生一坚固的脊骨突。触角上共发现7类11种感器, 分别为毛形感器、 刺形感器、 锥形感器（包括单孔形和多孔形）、 多孔板形感器（包括长形和圆形）、 腔锥形感器（分为3种类型）、 栓锥形乳突状感器、 角锥形感器。结合表面特征和内部结构, 锥形感器、 多孔板形感器、 栓锥形乳突状感器和腔锥形感器类型1为有孔型, 为化学感器; 无孔型的毛形感器和刺形感器是机械感器, 但腔锥形感器类型2和3为本体感器或湿热压力感器; 最为特异的为角锥形感器, 其厚壁无孔, 逆向触角主轴, 为该科昆虫所特有, 推测可防止传粉榕小蜂进入榕果时滑脱。这些结果将有助于理解木瓜榕传粉榕小蜂特异性行为, 并为下一步开展电生理研究, 揭示其信息化学物质利用和分配模式奠定基础。     

Electrophysiological responses to sucrose from a gustatory sensillum on the larval maxillary palp of the spruce budworm,Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)

The tip of the maxillary palp in the spruce budworm and other lepidoptera larvae contains a cluster of eight basiconic sensilla. Except for a few electrophysiological recordings from the entire group of these sensilla, no information is available on the response characteristics of any individual one. Using a compound microscope at 320x magnification, with a long working-distance objective, it has been possible to view individual sensilla and to record electrical responses from three of these to some gustatory stimuli in preliminary tests. One of them, sensillum (L1), contains a sugar-sensitive neuron whose responses to a range of sucrose concentrations are reported here. The physiological characteristics of this neuron differ from those of the known sugar-sensitive neuron in the lateral styloconic sensillum on the galea of these same insects. Input of chemosensory information about the same gustatory stimulus by more than one neuron in different mouthpart sensilla is discussed in relation to the feeding habits of the spruce budworm.     

六斑月瓢虫触角及感觉器扫描电镜观察

[目的] 明确六斑月瓢虫雌雄成虫触角感觉器种类、分布及形态特征。[方法] 利用扫描电子显微镜对六斑月瓢虫雌、雄成虫触角形态及触角感受器超微结构进行观察。[结果] 六斑月瓢虫成虫触角由柄节、梗节和鞭节组成,柄节长度与宽度显著大于梗节长度与宽度;鞭节分为9个亚节,末端3节横向膨大呈锤状。雌雄成虫触角上共有8种感觉器:刺形感觉器(SC)、毛形感觉器(ST)、锥形感觉器(SB)、腔形感觉器(CaS)、钟形感觉器(CS)、哑铃形感觉器(DS)、香肠形感觉器(SS)及B?hm氏鬃毛感觉器(BB)。以毛形感觉器和刺形感觉器分布最广,遍布触角;B?hm氏鬃毛仅存在于触角柄节与梗节;触角鞭节第9亚节顶端密布7种触角感觉器。六斑月瓢虫雌雄成虫触角长度、触角感觉器类型及分布无显著差异。[结论] 六斑月瓢虫成虫触角上共有8种感觉器,其触角可能具有感知机械刺激、识别化学信息素及感受温湿度变化的作用。本研究为进一步了解六斑月瓢虫触角与其行为间的关系提供基础资料。     

External morphology of antennal sensilla of trichogramma australicum girault (Hymenoptera : Trichogrammatidae)

External morphology of antennal sensilla on female and male Trichogramma australicum (Hymenoptera : Trichogrammatidae) was examined using scanning electron microscopy. Antennae show strong sexual dimorphism in structure and types of sensilla. The female antenna displays 14 types of sensilla: basiconic capitate peg sensilla (types 1 and 2), campaniform sensilla, chaetica sensilla (types 1–3), coeloconic sensilla, falcate sensilla, placoid sensilla (types 1 and 2), styloconic sensilla and trichoid sensilla (types 1–3). The male antenna displays 12 types of sensilla: basiconic capitate peg sensilla (type 2), campaniform sensilla, chaetica sensilla (types 1–5), coeloconic sensilla, placoid sensilla (type 1), and trichoid sensilla (types 3–5). Falcate and styloconic sensilla occur only on the female antenna. Both sensilla probably are associated with host examination, host discrimination and oviposition behaviour. Male antennal trichoid sensilla types 4 and 5 are probably associated with courtship behaviour, because these types occur only on the male. We propose the term “falcate sensilla” for a unique female antennal sensilla; the number of falcate sensilla may be used for identification of Trichogramma spp. In addition, we report the presence of placoid sensilla type 2 and difference in structure of coeloconic sensilla in T. australicum. Variation in structure and position of antennal sensilla are discussed.     

Sensilla of the antennal flagellum in Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae)

Seven sensilla were observed on the antennal flagellum of both sexes of Pseudaletia unipuncta (Haw.) (Lepidoptera : Noctuidae): chaetica Type I and Type II), styloconica, coeloconica, trichodea (Type 1a, b, c and Type II a, b), auricillica, nonporous and squamiformia. The major sexual dimorphism noted was the presence of Type la trichodea on the male antennae, suggesting that this sensillum may contain the receptor sites for the female sex pheromone. Differences in the size of sensilla coeloconica, auricillica and trichodea (Types Ib, IIb) were noted between sexes, being larger in females. Adults obtained from larvae fed on artificial diet generally had longer antennae and sensilla than those fed on young oat plants.     

红火蚁触角及其上感受器的扫描电镜观察

应用扫描电镜对采自我国广东吴川和广西南宁的红火蚁Solenopsis invicta Buren工蚁、有翅雌蚁和雄蚁触角的形态和感受器类型进行了研究。结果表明,工蚁、雌蚁和雄蚁的触角存在着较大的差异。工蚁和雌蚁的触角为膝状,末端2或3节膨大为棒状,雌蚁的触角11节,比工蚁(10节)的多1节。雄蚁的触角为线状,12节,末端没有棒状结构。工蚁和雌蚁的触角上共有7种感受器,分别为毛形、曲毛形、锥形、腔锥形、坛形、刺形感受器和Böhm氏鬃毛,前6种感受器大多集中在触角末端的棒节,棒节以外的各鞭节上主要是毛形感受器,Böhm氏鬃毛则分布于柄节和梗节的基部,鞭节上无此感受器。雄蚁触角上也有7种感受器,但其分布与工蚁的有较大差异,Böhm氏鬃毛的分布与工蚁和雌蚁的相同,锥形感受器分布在末端2节,坛形感受器分布在末端3节,其余的4种感受器较为均匀地分布在各鞭节,而不是聚集在端节。另外,新发现雄蚁中存在一种与已知的毛型感受器不同的超短毛状感受器,定名为超短毛感受器。最后,对红火蚁的触角及感受器在蚁群社会性行为中的作用进行了讨论。     

Olfactory responses and sensilla morphology of the blackcurrant leaf midge Dasineura tetensi

Olfactory responses of Dasineura tetensi (Rubs) (Diptera; Cecidomyiidae) to leaf volatiles of blackcurrant (Ribes nigrum) were tested in a 4-way olfactometer. Newly emerged virgin females showed no response to the leaf volatiles emitted from a blackcurrant shoot. Newly emerged males (which are known to respond to a pheromone released by the female) also showed no response to the leaf volatiles. Two hours after mating females responded positively, indicating that leaf volatiles may play a role in host plant finding. Scanning (SEM) and transmission (TEM) electron microscopy of the antennae of D. tetensi showed that males and females share five sensillum types; sensilla chaetica (mechanoreceptors), sensilla trichodea, sensilla basiconica, uniporous peg sensilla and circumfila (chemoreceptors). The sensilla chaetica and sensilla trichodea resemble those found on other insects. Sensilla basiconica were found on all antennal subsegments except the tip. These are multiporous receptors with five unbranched dendrites filling the lumen. Small peg sensilla located on the tips of both male and female antennae may function as contact-chemoreceptors. The circumfila, which are a unique type of sensilla found only on cecidomyiid antennae, form loops around each of the antennal subsegments, being attached to the surface by a series of stalks. TEM revealed that each stalk consisted of one sensillum containing a single highly branched dendrite. The distal regions of the walls of each sensillum are fused together to form the circumfila. Circumfila have multiporous walls and a lumen filled with multiple branches of dendrites. Their structure suggests that they are important olfactory receptors in both the male and female.