Labial tip sensilla of Blissus leucopterus leucopterus (Hemiptera: Blissidae): Ultrastructure and behavior

The cuticular sensory receptors that are found on the apex of the labium of hemipterans play an important role in their feeding behavior. In this study we describe the ultrastructure, number, and distribution of sensilla on the labium apex of the chinch bug, Blissus leucopterus leucopterus. Each apical field of sensilla on the labium contains 11 uniporous peg sensilla and one sensillum chaeticum. The uniporous peg sensilla are innervated by 4–5 bipolar neurons that send dendrites in the lumen of each peg. Three neurons are associated with each sensillum chaeticum, two neurons have dendrites in the lumen of the sensillum, and the third dendrite ends in a tubular body at the base of the sensillum. Behavioral tests that involve chemical blockage of the sensory receptors show the importance of the labial sensilla in feeding behavior. Both morphological and behavioral evidence indicate that the labial sensilla have a chemosensitive function.     

Morphology and distribution of antennal sense organs and diversity of mouthpart structures in Odontopus nigricornis (Stall) and Nezara viridula L. (Hemiptera)

Sensilla on the antennae and labial tip of the adult hemipteran bugs, Odontopus nigricornis Stall (Pyrrhocoridae) and Nezara viridula L. (Pentatomidae) (Hemiptera) were examined with an electron microscope in order to study their morphology, distribution, and possible chemo- and mechanoreceptive functions for food detection. The antenna contains 2 types of sensilla trichodea and 3 types of sensilla basiconica on the flagellar segments. S. trichodea are most abundant in both insects. Long slender hairs with elevated bases, and slightly shorter hairs with flexible sockets at the bases are common in both species. However, on each antennal segment of O.nigricornis, 1 or 2 rows of very long trichodeal hairs with longitudinal grooves and blunt tips are visible. At the periphery, S. basiconica with bulbous bases were also observed in addition to sensilla chaetica. The tripartite apex of the labium in both species consists of 2 lateral lobes and an apical plate. Each lateral lobe possesses a field of 11 thick-walled uniporous peg sensilla and long non-porous hair sensilla. Mandibular stylets of O.nigricornis have barbs, which may help in penetrating the hard seed coat. These insects made circular holes while feeding on the seed. A possible mechanism of feeding is discussed.     

Sensory receptors on the adult labial and maxillary palpi and galea of Cicindela sexguttata (Coleoptera: Cicindelidae)

Five types of sensilla are situated on the apical area of the labial and maxillary palpi and galea of Cicidela sexguttata. Large, conical, and peg-like sensilla are in rows on the central region of each palpus. These sensilla have a hollow cuticular peg, with an apical pore and multi-innervation. This central region of palpal sensilla is surrounded by campaniform sensilla that are disc-shaped and small conical peg sensilla. A similar type of conical sensillum as the found in the palpal central region is situated around the periphery of the palpal apex and apex of the galea. This conical peg sensillum is located in a shallow depression and is structurally similar to the other peg sensilla, but it has a mechanoreceptor neuron attached to the cuticular base of the sensillum. A long, single, trichoid sensillum is situated in the center of the galea and is hollow, thick-walled, porous, and multi-innervated. The apices of the palpi and galea have a large number of dermal gland openings that actively secrete a substance during the feeding process of the tiger beetle. © 1995 Wiley-Liss, Inc.     

Sensilla on the antennae,mouthparts, and body of the larva of the alfalfa weevil,hypera postica (gyllenhal) (Coleoptera : Curculionidae)

The morphology of the antennal complex, major mouthpart sensilla and body sensilla of alfalfa weevil larvae, Hypera postica (Gyllenhal) (Coleoptera : Curculionidae), is described. All instars have a short 1-segmented antenna with a large, multiporous sensillum basiconicum (s.b.) on its apex. Surrounding the s.b. are 2 unusual sensilla auricularia, and 3 small s.b., 2 of which are uniporous. The galea of the maxilla contains 11 s.b. and the apex of the palpus bears 12 thick-walled, uniporous s.b. The labial palpus has 10 similar s.b. apically.First-instar larvae have long, rodlike sensilla with a large, thick-walled bulbous apex; they are probably mechanoreceptors. The other 3 instars bear unusual nodulated and ridged sensilla with thin walls and a fragment-filled lumen. Sensilla on the apex of the thoracic legs and prolegs are also illustrated.     

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.     

Contact chemoreceptors among wind-sensitive head hairs of Locusta migratoria L. (Orthoptera: Acrididae)

The peg sensillum, a type of sensillum intermingled with the long hair sensilla in the hair fields on the head of Locusta migratoria (Orthoptera: Acrididae), was studied by light, transmission and scanning electron microscopy. The peg sensilla have the features typical of contact chemoreceptors; each peg is innervated by 5 sense cells; and 4 of the dendrites, enclosed within a “dendritic canal”, pass through the central lumen of the peg to the distal part, below the apical pore. The 5th dendrite ends in a tubular body at the caudal side of the peg articulation. Each distal segment of the 5 dendrites has a ciliary structure (9 ×2+0) at the transition to the short proximal segment, which in each case turns to the side to merge with the soma of the bipolar sense cell. Four sheath cells are associated with the group of sense cells and they are surrounded by a 5th, special epidermis cell. The innermost (thecogen) sheath cell (No. 1) encloses the receptor-lymph space 1 and forms the cuticular sheath; sheath cells 2 and 4 form the boundary of the large volume of receptorlymph space 2. The number of sheath cells is discussed with reference to other insect sensilla and in a phylogenetic context.     

Sensilla and Cuticular Appendages on the Female Abdomen of Lasioptera rubi (Schrank) (Diptera,Cecidomyiidae)

Studies by SEM and TEM revealed 6 types of integumental appendages on female uromeres VIII-X in Lasioptera rubi: microtrichia, not innervated; spines, probably without sensory function; nonporous sensory hairs, each containing one dendrite ending with a tubular body indicating a tactile function; uniporous sensory hairs, each innervated partly by 3 dendrites indicating a chemosensory function, partly by an additional dendrite with a tubular body indicating a tactile function; scoop-like sensilla, each containing partly a branched structure of dendrites in the distal half of the sensillum indicating an olfactory function, partly an unbranched dendrite ending at a pore near the base of the sensillum, most probably registrating chemical stimuli by contact or gustation; finally, nonporous bristles, all or some of them innervated, in a manner indicating a tactile function. In addition, two scolopophorous proprioceptors were found inside uromere X. The nonporous sensory hairs, the uniporous sensory hairs and the scolopophores may be used by the midge to determine the mechanical and chemical properties of potential oviposition sites. The spines and nonporous bristles may function as conidia carriers.     

Ultrastructure of sensilla of the female mountain pine beetle,Dendroctonus ponderosae hopkins (Coleoptera : scolytidae)

The objective of this study is to provide a morphological foundation for electrophysiological studies on the sensilla of D. ponderosae. The sensilla on the antennae, labial and maxillary palps, galeae and fore-tarsi are described. The antennal club has several types of sensilla: (1) multiporous non-socketed pegs of varying lengths innervated by 2 neurons, (2) uniporous socketed pegs innervated by 5 neurons, (3) pegs innervated by only 1 neuron at the base, (4) tapered uniporous cones innervated by 4 neurons, and (5) fluted multiporous, non-socketed cones innervated by 4 neurons. All hairs on the fore-tarsi are innervated by 1 neuron at the base. The labial and maxillary palp-tips have: (1) digitiform organs, (2) campaniform organs, (3) uniporous, socketed pegs innervated by 5 neurons, and (4) non-porous pegs innervated by 3 neurons. Two small, uniporous, socketed pegs innervated by 4 neurons, are set into the maxillary palp-tip sidewall, and a single, sharp-tipped peg, innervated by 1 neuron, is found on the tip. These 2 types are not seen on the labial palps. The ultrastructure of each type of sensillum found on both the maxillary and labial palps is similar; however, the number of uniporous and non-porous pegs on the labial palps is about half those of the maxilla. The maxillary galeae each have 3 types of sensilla: (1) numerous pegs of various sizes innervated by 1 neuron, (2) a single, possibly uniporous socketed peg innervated by 5 neurons, and (3) another single non-porous non-socketed peg innervated by 2 neurons.     

Antennal sensilla of the adult alfalfa weevil,Hypera postica (Gyllenhal) (Coleoptera : Curculionidae)

Sensilla on the antennae of adult alfalfa weevils, Hypera postica (Gyllenhal),were studied with scanning and transmission electron microscopes to determine the sensilla structure and possible chemo- and mechanoreceptive functions for food detection.Five types of sensilla trichodea (s.t.) occur (length 35 – 135 μm); s.t. I (71 – 135-μm long)are thick-walled without radial pores and s.t. II (50 – 75-μm long), the most abundant s.t., are multiporous. Bifurcated and trifurcated types (75 –85 -μm long) also occur. Five types of sensilla basiconica (s.b.) are present and include s.b. I (26 – 50-μm long), a fluted uniporous type with a rugose surface and encircling ledges; s.b. II (15 – 35-gm long), a common multiporous type with many longitudinal grooves; and s.b. IV (4 – 7-μm long), which are 3 apical, short pegs. Sensilla styloconica occur in a small band among the s.b. type II. One or 2 epidermal gland ducts are adjacent to most s.t. and some s.b.     

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.     

Structure of galeal sensory complex in adults of the red turnip beetle,Entomoscelis americana brown (Coleoptera,Chrysomelidae)

Summary The internal and external structure of the galeae of the adult red turnip beetle, Entomoscelis americana, was studied using SEM and TEM. The galea broadens from base to truncated tip and its sides are of thick, sculpted cuticle invested with pores and coarse spines. The tip is of thinner, flexible cuticle covered with 8–12 uniporous, blunt-tipped apical pegs and a single, aporous, sharply-pointed apical hair.The coarse spines are singly innervated probable mechanosensilla owing to the tubular body at the distal end of the dendrite. These sensilla likely act as tactile hairs monitoring galeal-effected movements of food particles into the functional mouth. The pores are associated with glands within the galea. The function of the presumed secretion is not known but may be to keep objects and dried saliva from sticking to the mouthparts.The apical pegs are innervated by five neurons, each producing a single dendrite. Four dendrites enter the single peg lumen and communicate with the terminal pore. The fifth differentiates into a tubular body that inserts into the peg base. These are typical insect contact chemosensilla that, because of their location, would taste incoming food.The apical hair has no pores but is innervated by two neurons, each extending a dendrite into the hair lumen in chemosensillar fashion. The sensory mode of this sensillum is unknown but is probably not mechanoor chemoreception. Many of its features, reminiscent of taste hairs, lead us to hypothesize that it represents a one-time chemosensillum recently modified to a new form and sensory mode.Because larval and adult E. americana share similar food plant requirements, we hypothesize that similarities will be seen in their mouthpart sensilla. Comparisons of the adults and larvae show the common features between their respective galeal taste hairs are only those of insect contact chemosensilla in general. However, the adult apical hair and the larval medial sensillum show striking specific structural similarities. We propose that these are true structural and functional homologues.     

Antennal sensilla of Magicicada cassini (fisher) (Homoptera : Cicadidae): Fine structure and electrophysiological evidence for olfaction

The antennae of Magicicada cassini (Homoptera : Cicadidae) (3–4 mm long) look similar in both sexes and consist of scape, pedicel, and a 5-segmented flagellum. The length of flagellar segment 1 varies independently in relation to head size and is slightly longer in females (0.96 mm) than in males (0.89 mm). The ventral side of flagellar segment 1 is covered with sensilla coeloconica comprising about 60 large, 10 medium-sized, and 35 small sensilla with pit diameters of 8–24, 6–10, and 2 μm, respectively. The large and the medium-sized sensilla coeloconica are multiporous single-walled sensilla with pore tubules, containing branched entangled dendrites from 3 receptor cells. The small sensilla coeloconica, situated primarily at the outer border of the sensillum field, are no-pore sensilla with inflexible sockets. They contain 2 unbranched dendrites extending to the tip of the peg, and 1 dendrite reaching to its base and wrapping around the other 2 dendrites. Small sensilla campaniformia (cap diameter 3 μm) are aligned at the outer border of the sensillum field and continue all along the flagellum. Up to 3 olfactory receptor cells were distinguished on the basis of their nerve impulse amplitudes through extracellular electrophysiological recordings from sensilla coeloconica, presumably large ones. They respond to stimulation by cyclic terpenoids with different but highly overlapping reaction spectra, and react selectively to structural variations of the molecules. No responses to CO₂, temperature or moisture were recorded.     

Ultrastructure of the antennal sensilla of the cockroach-egg parasitoid,Tetrastichus hagenowii (Hymenoptera: Eulophidae)

Eggs of a number of cockroach species are parasitized by Tetrastichus hagenowii. The ultrastructure of the sensilla on the antennae of females and males was examined by scanning and transmission electron microscopy. The females have two types of multiporous plate sensilla while the males have only one. Type 1 is found in females and males and has a relatively thin cuticular wall and many pores, while type 2 is found only in females and has a relatively thick cuticular wall and few pores. Both sexes have nonporous, thick-walled, socketed hairs; multiporous, nonsocketed hairs; multiporous, thick-walled pegs; and terminal hairs. In addition, males have multiporous, nonsocketed, long hairs. The sensilla are similar, in many respects, to the sensilla of other chalcid parasitoids. The antennal sensilla of female T. hagenowii are probably involved in ovipositional behavior. The multiporous, long hairs of the male possibly receive stimuli during mating behavior A chemoreceptive function is proposed for the multiporous plate sensilla.     

Antennal sensilla in five Psychodini moth flies (Diptera: Psychodidae: Psychodinae) pollinators of Arum spp. (Araceae)

The pollination of the genus Arum (Araceae) is mainly achieved by deception, the floral odour mimicking the pollinator ovipositing site. In order to discover the sensory organs involved in this attraction, we have studied the antennae of five species of psychodine moth-flies (former Psychoda sensu lato=Psychodini), pollinators of Arum spp. The antennae of the five Psychodini reveal seven types of sensilla: multiporous tribranched sensilla basiconica (sensilla ascoidea), multiporous sensilla basiconica, multiporous sensilla coeloconica, multiporous sensilla auricillica, uniporous sensilla basiconica, aporous sensilla chaetica, aporous Böhm’s sensilla. Each species possesses three, five or six of these sensillum types. All the multiporous sensilla are probably olfactory receptors while the uniporous sensilla basiconica must possess a contact chemoreceptive function. The multiporous tribranched sensilla basiconica (s. ascoidea), present in all the species, are the best candidates for the reception of the odours given off by the ovipositing sites and the inflorescences of Arum. The multiporous sensilla basiconica and the multiporous sensilla coeloconica may be involved respectively as CO₂ receptors or thermoreceptors. Psychoda phalaenoides, which is the main pollinator of A. maculatum, is the species which possesses the largest number of antennal sensilla. The sexual dimorphism, studied only in Psycha grisescens, as concerns the number of sensilla and the absence of a sensillum type which differ according to sex, is difficult to interpret.     

Fine structure of antennal putative thermo-/hygrosensilla of adult Rhodnius prolixus Stål (Hemiptera: Reduviidae)

At least five nonporous sensilla with inflexible sockets (npsensilla) occur on each antenna of both sexes of adult Rhodnius prolixus. Externally the sensillum appears as a short, rounded peg set into a pit surrounded by a depression. A very electron-dense material occurs in the peg lumen and the inner aspect of the pit. Filamentous extensions of this material radiate into the overlying outlets. Each sensillum is innervated by three neurons with unbranched dendrites. Two dendrites extend to the peg tip and distally are covered by a dendritic sheath. The portion of these dendrites within the sheath contains a large number of microtubules. The third dendrite terminates near the base of the dentritic sheath and partially wraps around the other two dendrites. Three sheath cells are associated with each sensillum. Based on similarities in structure with sensilla of known function it is probable that the np-sensilla of R. prolixus are thermo-/hygrosensilla responding to cold, dryness and wetness. The sensilla have a number of structural similarities with insect rectal sheath cells known to absorb atmospheric water by electroosmosis. Possibly this process leads to volumetric alterations of cuticular elements associated with the dendrites and ultimately to mechanotransduction.     

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.     

External morphology of sensilla in the sacculus of an antennal flagellum of the fruit fly Drosophila melanogaster Meigen (Diptera : Drosophilidae)

Sensilla in the sacculus of an antennal 3rd segment, a funiculus, of the fruit fly, Drosophila melanogaster (Diptera : Drosophilidae) have been examined by scanning electron microscopy. The sacculus was divided into 3 cavities in its interior. A morphologically distinct group of sensilla was present in each cavity. Grooved sensillum (GS), found in the largest cavity, was further subclassified on the basis of the side wall sculpture into 3 subgroups: GS Ia, GS Ib and GS II. GS Ia was 4 μm long and had 10–12 grooves (0.25 μm wide) and GS Ib was 3.8 μm long and had 6–9 grooves (0.25-0.4 μm wide). GS 1a and GS Ib were inferred to be olfactory and thermoreceptive, and olfactory, respectively. GS II was 3.2 μm long, and had 4–5 grooves. Basiconic sensillum (BS), found in the smallest cavity, was 4.5 μm long and had an irregularly sculptured side wall, suggesting the presence of numerous irregular-shaped olfactory pores. Blunt-tipped sensillum (BTS), found in the middle-sized cavity, was 1.9 μm long and had a smooth-surfaced side wall and a button-like structure on its apex. These features suggested that BTS was hygro- and thermoreceptive.     

Fine structure of the galeal styloconic sensilla of larval Lymantria dispar (Lepidoptera: Lymantriidae)

Lepidopteran larvae possess two pairs of styloconic sensilla located on the maxillary galea. These sensilla, namely the lateral and medial styloconic sensilla, are each comprised of a smaller cone, which is inserted into a style. They are thought to play an important role in host-plant selection and are the main organs involved in feeding. Ultrastructural examination of these sensilla of fifth instar Lymantria dispar (L.) larvae reveal that they are each approximately 70 um in length and 30 um in width. Each sensillum consists of a single sensory peg inserted into the socket of a large style. Each peg bears a slightly subapical terminal pore averaging 317 nm in lateral and 179 nm in medial sensilla. Each sensillum houses five bipolar neurons. The proximal dendritic segment of each neuron gives rise to an unbranched distal dendritic segment. Four of these dendrites terminate near the tip of the sensillum below the pore and bear ultrastructural features consistent with contact chemosensilla. The fifth distal dendrite terminates near the base of the peg and bears ultrastructural features consistent with mechanosensilla. Thus, these sensilla each bear a bimodal chemo-mechanosensory function. The distal dendrites lie within the dendritic channel and are enclosed by a dendritic sheath. The intermediate and outer sheath cells enclose a large sensillar sinus, whereas the smaller ciliary sinus is enclosed by the inner cell. The neurons are ensheathed successively by the inner, intermediate, and outer sheath cells.     

Ultrastructural Analysis of the Sensilla of Austroperipatus aequabilis Reid, 1996 (Onychophora,Peripatopsidae)

The head of Austroperipatus aequabilis bears five types of sensilla. which were examined by electron microscopy. They differ from each other in position, shape of outer sensory elements and cuticular socket structures. Thus, we distinguish sensilla with sensory hairs, sensilla with sensory bulbs, cone-shaped sensilla. sensilla with sensory bristles, and sensilla of the lips. They are composed of up to 15 cells, which can he separated into four cell types. The most frequent cell type is the bipolar receptor cell that occurs in all sensilla. The apical surface of this primary receptor cell is characterized by one or two partly branched cilia with a basal 9 × 2 + 0 pattern of microtubules. A modified bipolar receptor cell was found in all sensilla bearing a sensory peg except for the sensilla equipped with sensory bristles. The apical dendrite extends to a long pale process which exclusively contains mitochondria and single microtubules. In all sensilla examined in this study at least one supporting cell occurs which is characterized by parallel microvilli. An additional function of this cell type as a part of the stimulus-conducting system is possible. In the sensillum with a sensory bulb two kinds of supporting cells occur. A unique cell type with an upside down position has regularly been found in all sensilla bearing a sensory peg. Apart from the sensilla they also occur within the labial epidermis. Since most sensilla contain several different receptor cells, they can be considered as complex sense organs. © 1998 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd. All rights reserved     

Morphology of antennal sensilla of the brown spruce longhorn beetle,Tetropium fuscum (Fabr.) (Coleoptera: Cerambycidae)

The antennal sensilla of the brown spruce longhorn beetle, Tetropium fuscum (Fabr.) (Coleoptera: Cerambycidae) were examined with particular focus on the sensilla present on the apical flagellomere. T. fuscum antennae are composed of 11 segments, namely the scape, pedicel, and nine flagellomeres. Nine types of sensilla were observed: three types of sensilla chaetica, sensilla trichodea, two types of sensilla basiconica, grooved peg sensilla, thick-walled sensilla, and Böhm bristles. Seven of these types were present on the apical flagellomere, the exceptions were sensilla chaetica type 3 and Böhm bristles. There were no significant differences in the distribution or density of sensilla present on the ninth flagellomere of males and females, except that males had significantly more sensilla chaetica type 1, which are put forward as the putative contact chemoreceptors for T. fuscum.