Morphology and distribution of the external labial sensilla in Fulgoromorpha (Insecta: Hemiptera)

The present paper describes the sensory structures on the apical segment of the labium in fifteen fulgoromorphan families (Hemiptera: Fulgoromorpha), using the scanning electron microscope. Thirteen morphologically distinct types of sensilla are identified: five types of multiporous sensilla, four types of uniporous sensilla and four types of nonporous sensilla. Three subapical sensory organ types are also recognized, formed from one to several sensilla, each characteristic of a family group. Sensilla chaetica (mechanoreceptive sensilla) fall into three categories dependent on length and are numerous and evenly distributed on the surface of the labium except where they occur on specialized sensory fields. The planthopper morphological ground plan is represented by two apical pair of sensory fields (dorsal and ventral) on which 11 dorsal pairs of sensilla (10 peg-like pairs + 1 specialized pair dome or cupola-like) and 2 ventral pairs of sensilla basiconica occur. Two main patterns (cixiid and issid) together with more specialized ones (derbid, lophopid, flatid and fulgorid) are reported. Disparity and diversity of the sensory structures are analyzed from a taxonomic and functional perspective. A gustatory function is provided for several chemoreceptive labial sensilla, as in the antennal flagellum sensilla in some other Hemiptera. This represents a more recently evolved function for the planthopper labium. Finally, further lines of study are suggested for future work on the phylogeny of the group based on the studied characters.     

The fowl tick, <Emphasis Type="Italic">Argas (Persicargas) persicus</Emphasis> (Ixodoidea: Argasidae): Description of the egg and redescription of the larva by Scanning Electron Microscopy

This scanning electron microscopy study revealed that the egg of Argas persicus was covered with chorion which appeared as a wrinkled layer containing regions of three textures. The first had elevated parts of slightly rough surface. The second had irregular smooth elevations; each carried numerous parallel horizontal foldings with vertical ridges. The last region had rough surface with irregularly shaped projections. Following the removal of the chorion, shell was observed to have one polar micropyle and numerous slit like openings. Length (L), width (W) and L/W ratio of the egg were measured. Investigation of larvae revealed extensively folded integument of idiosoma and spherical or elongated tubercles on dorsal plate. Mouth enclosed between ventral hypostome and two dorsal chelicerae. Hypostome carried four longitudinal rows of conical denticles. Each chelicera was made up of two segments; the basal one appeared as a pocket for the distal one. Haller’s organ consisted of an anterior pit containing seven sensilla and a posterior capsule with four apertures. Distribution of chemo- and mechano-sensilla on the body was examined. Measurements of whole body, idiosoma, dorsal plate, capitulum, hypostome, palp and different types of sensilla both on the body and Haller’s organ are also presented.     

Microscopic anatomy of male tegumental glands and associated cuticular structures in Titanethes albus (Crustacea: Isopoda)

Male glandular organs characterized by porous surfaces with hair-like cuticular elaborations are known from several trichoniscid isopods. In the subterranean species Titanethes albus, males possess paired tubercles with numerous hairs and pores dorsally on the pleon. We analyzed the microscopic anatomy of these structures with scanning and transmission electron microscopy. Diverse epicuticular formations and numerous sensilla, which are probably chemoreceptive, are present on the tubercles. We found several secretory surfaces on the pleon in addition to the dorsal tubercles. We also examined the distribution, architecture and ultrastructure of male-specific glands in T. albus with light and transmission electron microscopy. Three distinct types of male-specific rosette glands are present in different parts of the pleon and in the uropods. Glands secreting on the dorsal tubercles contain stellar central cells. The ultrastructure and histochemical staining properties of male-specific glands in T. albus suggest that they produce peptides which might function as contact pheromones.     

The sensory equipment of a spider – A morphological survey of different types of sensillum in both sexes of Argiope bruennichi (Araneae,Araneidae)

Spiders show a wide range of sensory capabilities as evidenced by behavioural observations. Accordingly, spiders possess diverse sensory structures like mechano-, hygro-, thermo- or chemoreceptive sensilla. As to chemoreceptive structures, only trichoid tip-pore sensilla were found so far that were tested for gustation. That spiders are also able to receive airborne signals is corroborated by numerous behavioural experiments but the responsible structures have not been determined yet. Here, we provide sensilla distribution maps of pedipalps and walking legs of both sexes of the wasp spider Argiope bruennichi whose biology and mating system is well explored. By means of scanning electron microscopy, we scrutinized whether there is in fact only one type of trichoid pore sensillum and if so, if there are deviations in the outer structure of the tip-pore sensilla depending on their position on the body. We also describe the external structure and distribution of slit sense organs, trichobothria and tarsal organs. Our study shows that all four sensillum types occur on pedipalps and walking legs of both sexes. As to chemosensory organs, only tip-pore sensilla were found, suggesting that this sensillum type is used for both gustation and olfaction. The highest numbers of tip-pore sensilla were observed on metatarsi and tarsi of the first two walking legs. Mechanosensitive slit sense organs occur as single slit sensilla in rows along all podomers or as lyriform organs next to the joints. The mechanosensitive trichobothria occur on the basal part of tibiae and metatarsi. Tarsal organs occur on the dorsal side of all tarsi and the male cymbium. The distribution maps of the sensilla are the starting point for further exploration of internal, morphological differences of the sensilla from different regions on the body. Cryptic anatomical differences might be linked to functional differences that can be explored in combination with electrophysiological analyses. Consequently, the maps will help to elucidate the sensory world of spiders.     

The formation of the epicuticle and associated structures inOniscus asellus (Crustacea,Isopoda)

Summary The integument of the woodlouse,Oniscus asellus, consists of a two-layered epicuticle, a largely lamellate procuticle — itself divided into two regions (pre-and postecdysial cuticles), and the epidermis. At the initiation of new cuticle production the epidermal cells become vacuolated and retract away from the cuticle. Apolysis occurs immediately after the cessation of postecdysial cuticle production. The formation of the epicuticle is unique among the arthropods since material aggregates along the distal epidermal membrane. By indenting, doubling back on itself, and incorporating septa, the epicuticle forms surface structures such as plaques and tricorns.The innervation, and so the receptive function of the tricorns is confirmed, but since there is no connection between the old and new receptors during premoult, sensory information from these exoreceptors must be severely curtailed. This may explain the biphasic moult in all isopods since it ensures that only half the body experiences this sensory deprivation at any one time. In terrestrial species there is the additional advantage of restricting the area of permeable new cuticle. The frequency of moulting may be due to the need to renew disrupted receptor surfaces.Tricorns do not appear to be the mechanoreceptors involved in the marked thigmotactic response of woodlice since they do not have the typical internal structure of such receptors; rather, the dendrite —which extends into the lumen of the tricorn —is protected from deformation by the previously unreported combination of a dendritic sheath and a cuticular tube. The modality of tricorns is possibly one of hygro-perception. One of the behavioural responses of woodlice to desiccation is aggregation. The numerical distribution of tricorns over the body surface is admirably suited to assist in the formation and maintenance of such aggregates during desiccation and to their observed dispersal when the relative humidity rises.     

Review of the Halieutichthys aculeatus species complex (Lophiiformes: Ogcocephalidae), with descriptions of two new species

The Halieutichthys aculeatus species complex is reviewed. Members of this clade are distinguished by the presence of tubercles on the tail and a reticulate dorsal pigmentation pattern. Three species are recognized, including two species new to science. A neotype is chosen for H. aculeatus. Halieutichthys bispinosus n. sp. is characterized by having relatively strong tubercles on the dorsal surface, a row of tubercles almost always present dorsal to the orbit, both sphenotic tubercles well developed and sharp, trifid principal tubercles on the disk margin with anterior spinelet enlarged, dense arrangement of tubercles on the tail and a comparatively large adult body size. Halieutichthys intermedius n. sp. can be distinguished from congeners by having both sphenotic tubercles strongly reduced, the tubercles almost always absent dorsal to the orbit, principal tubercles on the disk margin uniformly short and blunt tubercles, with all spinelets generally reduced.     

Unculi (Horny Projections Arising from Single Cells), an Adaptive Feature of the Epidermis of Ostariophysan Fishes

Horny projections arising from single cells, or unculi, of ostariophysan fishes are described and discussed by several workers, but they occur in many more families and genera than previously reported and their status as a key adaptive feature has not been recognized. Unculi are related morphologically to the relatively well-known multicellular horny tubercles (including breeding tubercles) of Ostariophysi and other fishes, but differ from them in anatomical distribution and function. Unculi are especially prominent features of portions of the epidermis in Cyprinoidei (carps, loaches, and their allies) and Siluroidei (catfishes); they also occur in Characoidei (characins) and Chanoidei (gonorynchs) but have not been found in any non-ostariophysan fishes. Unculi evidently are functionally significant on the lips and other mouth parts (especially in groups lacking jaw teeth), on the ventral surface of the paired fins in many bottom-dwellers, and on several neomorphic features including epidermal plaques and tubercles in various catfishes, the thoracic adhesive disc of sisorid catfishes, and the mental adhesive disc of the cyprinid genus Garra . In some sisorid and akysid catfishes virtually the entire external body surface is covered by unculiferous plaques or tubercles. Possible functions of unculi include: mechanical protection of the skin, rasping, adhesion, and hydrodynamic effects. Unculi seem to have been especially important in the diversification of cyprinoid feeding habits and in the adaptation of bottom-dwelling cyprinoids and siluroids to swift-water habitats, especially in Asia.     

Minute tubercles on the skin surface of larvae in the Korean endemic bitterling, Rhodeus pseudosericeus (Pisces, Cyprinidae)

The morphology and distribution of the minute tubercles on the skin surface of larvae in Korean bitterling, Rhodeus pseudosericeus, were observed during larval development. Just after hatching, the epidermis of the larvae consists of a thin single cell layer having smaller basophilic flat or round‐flattened basal cells. As the larvae grow, the epidermis contains more small flat cells and large epidermal cells that are round or hemisphere‐shaped. These large unicellular epidermal cells, called minute tubercles, consist of more or less homogeneous cytoplasm that is PAS (Periodic acid‐Schiff method) positive. They are more densely distributed in the wing‐like yolk sac projection. Vestigial minute tubercles occur in the body region and the caudal fin‐fold region. These minute tubercles grow in number and height from 6 to 8 days after hatching onward. However, they become reduced in height and number as the larvae develop. At day 31 after hatching (i.e. free‐swimming stage), minute tubercles no longer exist on the larval skin. The sequence of occurrence and gradual disappearance of these cell structures are described and histologically documented for comparative purposes of beta, taxnomomic and environmental studies.     

Fine structure and primary sensory projections of sensilla on the maxillary palp of Drosophila melanogaster Meigen (Diptera : Drosophilidae)

Three types of hairs were identified on the maxillary palp of Drosophila melanogaster Meigen (Diptera : Drosophilidae): (i) single-walled, multiporous sensilla basiconica, which constitute 75% of the innervated hairs; (ii) thick walled non-porous sensilla trichodea, which make up the remaining 25% of the innervated hairs; and (iii) numerous spinules, which are un-innervated. These sensilla basiconica uniformly contain 2 bipolar sense cells, whereas sensilla trichodea have a single dendrite with a tubular body at the base of each hair. A majority of the sensilla basiconica is located on the distal half of the dorsal surface, whereas sensilla trichodea are positioned on the tip and entire ventrolateral ridge of the palp. Approximately 125 axons of the sense cells join to form a single nerve. The structure of sensilla basiconica and sensilla trichodea suggests that they are olfactory and mechanosensory respectively. The contact chemoreceptors (gustatory sensilla) are conspicuously absent on the maxillary palp.Golgi silver impregnations and cobalt fills show that the primary sensory fibres from sensilla trichodea and sensilla basiconica on the maxillary palp project in the posterior suboesophageal ganglion (SOG) and the antennal lobe respectively. A single fibre projects separately either in the SOG or in the antennal lobe. In the antennal lobe, the input received from sensilla basiconica is usually bilateral and at least 5 glomeruli are innervated symmetrically on either side from both the palps.This study suggests that the sensory neurons are capable of making selective projections in the specific regions of the brain. Accordingly, the fibres from a sensillum project to the brain with respect to their functions and the individual glomeruli represent functional units of the brain, receiving inputs in a characteristic combination.     

Morphology and ultrastructure of sense organs on the ovipositor of Trybliographa rapae,a parasitoid of the cabbage root fly

The ovipositor of the parasitoid wasp Trybliographa rapae was examined by scanning and transmission electron microscopy. Characteristic peg-like sensilla with a cuticular ring at the base are found at the tip of the ventral valves, where they occur in a characteristic arrangement of triplets. The unusual basal structure probably protects the sensilla against damage during movement through the substrate and piercing of the host cuticle. These sensilla are each innervated by six dendrites, some of which have lamellated tips, generally considered to be characteristic of thermosensitivity. It is suggested that the remaining dendrites are gustatory, and as such probably respond to factors present in host haemolymph. A second type of peg-like sensillum is found on both the dorsal and the ventral valves. These are set in deep pits so that only the tip of the peg protrudes above the surface of the cuticle. These occur along the length of the ovipositor shaft and ultrastructural studies reveal the pegs to be innervated by a single mechanosensitive dendrite, probably monitoring the movement of the ovipositor through the substrate.     

Reproductive strategies and parasitization behavior of Ageniaspis citricola, a parasitoid of the citrus leafminer Phyllocnistis citrella

We describe the number, distribution, and function of sensilla located on different organs of Lobesia botrana (Lepidoptera: Tortricidae) females using scanning electron microscopy, selective staining, and contact electrophysiology. The tarsi of the prothoracic legs bear contact chemo‐mechanoreceptor sensilla chaetica (5–13 per tarsomere), arranged in rings mainly concentrated on ventral surfaces, and different mechanosensory structures (sensilla chaetica, sensilla squamiformia, sensilla campaniformia, and spines). A single contact chemo‐mechanoreceptor sensillum chaeticum is present between the claws on the pretarsus. The ventral surface of the ovipositor lobes is covered with numerous mechanosensory sensilla chaetica of different types, out of which 10 have a contact chemosensory function. Putative contact chemo‐mechanoreceptor sensilla were also observed on the proboscis and antenna. Longitudinal rows of alternated sensilla styloconica and basiconica are present on the distal part of the proboscis, and rings of sensilla chaetica are present at the antennal tip. The sensilla on these body parts may play different roles in the selection of an oviposition site.     

Ultrastructure of tarsal sensilla and other integument structures of two Pseudocellus species (Ricinulei, Arachnida)

Ricinuleids are one of the least investigated groups of Arachnida. In particular, knowledge of their ultrastructure is poor. Observations of the distal tarsomeres of ricinuleids show differences in their shape and equipment of surface structures. Legs I and II are used by the Ricinulei to explore their surroundings with tentative movements. The tarsomeres of these legs show similarities in shape and surface structures that distinguish them from those of legs III and IV. In this study, 11 different structures of the tarsomere surfaces of two cave-dwelling species, Pseudocellus pearsei and P. boneti from México, were investigated for the first time with scanning and transmission electron microscopy and discussed regarding their possible function: 1) a single treelike ramifying seta resembles a no pore single-walled (np-sw) sensillum; 2) setae occurring in a small number and possessing a bipartite shaft represent terminal pore single-walled (tp-sw) sensilla. The surface of the proximal half of the shaft shows small branches. The distal half has a smooth surface; 3) long setae with conspicuous longitudinal lamellae show characteristics of chemoreceptive wall pore single-walled (wp-sw) sensilla; 4) frequent small wp-sw sensilla with flat and irregular lamellae; 5) very short wp-sw sensilla occurring solitary or in groups; 6) a few short setae with smooth surface correspond to wp-sw sensilla; 7) a single short clubbed seta articulating in a flat pit is considered to be a np-sw sensillum; 8) common long setae with a pointed tip show characteristics of mechanoreceptive np-sw sensilla; 9) ventral setae with adhesive and mechanosensory function are accompanied by multicellular "class III" glands; 10) slit organs with mechanoreceptive function; and 11) dome-like tubercles with no indication of sensorial function. Several of these sensilla form a sensory field on the dorsofrontal surface which is particularly pronounced on the distal tarsomeres of legs I and II.     

Morphology of the antennal sensilla of the cabbage root fly,Delia radicum L. (Diptera : Anthomyiidae)

The antennal sensilla of Delia radicum L. (Diptera : Anthomyiidae) were studied by scanning electron microscopy. On the scape and pedicel, grooved socketed bristles and setiferous plaques were found. There are 4 types of surface sensilla on the funicle: trichoid, basiconic, clavate, and grooved. Their numbers and distribution are described. There are 3–4 single-chambered pits on the dorsal surface of the funicle of both sexes and these contain basiconic sensilla. On the ventral surface, there is one multi-chambered pit, which contains 5 types of sensilla: grooved s., smooth-walled conical s., smooth-walled tapered s., striated s. and a novel type, flattened sensilla. These results are compared with previously published studies on several other fly species.     

Morphology and histology of secretory setae in terrestrial larvae of biting midges of the genus Forcipomyia (Diptera: Ceratopogonidae)

Apneustic larvae of the genus Forcipomyia possess unique secretory setae located on the dorsal surface along the body in two rows, one pair on each thoracic and abdominal segment and two pairs on the head. Morphological and histological studies of secretory setae in fourth instar larvae of Forcipomyia nigra (Winnertz) and Forcipomyia nigrans Remm indicate they are modified mechanoreceptors (sensilla trichodea) in which the trichogen cell is a glandular cell producing a hygroscopic secretion. The cytoplasm of the glandular trichogen cell fills the lumen of a secretory seta, which shows one or more pores on the apex. The cytoplasm contains numerous microtubules responsible for transportation of proteinaceous vesicles, and an extremely large polyploid nucleus typical of gland cells. The main role of the hygroscopic secretion is to moist the body and thus facilitate cuticular respiration.     

The Anatomy of the Tarsi of Schistocerca gregaria Forskål

Summary The tarsus of S. gregaria is divided into three units (here called segments) and an arolium set between a pair of claws. The first segment bears three pairs of pulvilli in the fore and middle legs, and one pair and two single pulvilli in the hind legs. Segment two bears a pair of pulvilli, segment three one long pulvillus and the arolium a similar pad on the undersurface. The outer layers of the arolium pad differ from those of the pulvilli in possibly lacking an epicuticle and in having a layer of cuticle which, unlike the corresponding layer in the pulvilli, does not stain with protein stains. The claws and dorsal surfaces bear trichoid sensilla, basiconic sensilla and campaniform sensilla. Smaller basiconic sensilla and canal sensilla occur on the proximal part of the pulvilli, and basiconic sensilla on the arolium undersurface. Internally the cuticle is modified in the arolium and pulvilli so that rods of probably chitin and resilin are formed. This would impart flexibility to the undersurfaces whilst retaining some degree of rigidity which might prevent damage to the small and delicate sense organs on the pulvilli. The tip of the arolium is specialised for adhesion, and there are two large neurones internally which could conceivably monitor attachment or detachment of the tip. There are chordotonal organs in segment three, and several other large neurones throughout the tarsus, some of which are associated with the slings of tissue holding the apodeme in a ventral position. Gland cells occurring in the dorsal epidermis of the adult mature male are also briefly described.     

Surface topography of the tegument of adult Schistosoma nasale Rao, 1933 from Sri Lanka

Scanning electron microscopical studies of adult male and female Schistosoma nasale are reported. The tubercles on the dorsal and dorso-lateral surfaces of unpaired male S. nasale are devoid of spines. In paired male worms the tubercles on the dorsal surface are large and also are devoid of spines, but some tubercles on the dorso-lateral surface possess spines. Pit-like openings are visible on the surface of the smooth tubercles. The oral and ventral suckers on the male worm are well developed and are invested with spines, as are the gynaecophoric canal and flap. Ciliated sensory receptors are distributed over the surface of the male worm. The oral and ventral suckers of the female worm are much smaller than those of the male: spines occur on both suckers. The surface of the female is non-tuberculate and is thrown into transverse folds. Pit-like openings are visible at higher magnifications. The anterior end of the female is heavily invested in ciliated receptors, whereas the posterior end is heavily spined. The surface topography of S. nasale is discussed in relation to other species of Schistosoma.     

Responses of dorsal tricorn-type sensilla on Ligia exotica

• 1.1. In order to investigate the function of the dorsal tricorn-type sensilla on Ligia exotica, the morphology and distribution of the setae and neural responses to certain stimulus modalities were studied.
• 2.2. These foraminate sensilla are found to occur over the body surface, except for several appendages and the ventral carapace (sternite); the dorsal carapace (tergite) is covered by only this type of sensilla.
• 3.3. The tricorn-type sensilla located on the dorsal carapace responded to mechanical, gustatory and olfactory stimulation.
• 4.4. The function of the tricorn-type sensilla on the dorsal carapace was discussed.

     

The surface microstructure of marine and terrestrial isopoda (Crustacea,Peracarida)

Summary The microstructure of the surface of thirteen marine littoral and two terrestrial isopods was investigated by scanning electron microscopy. A great diversity of surface ornamentation is present, including non-sensory microscales, pits, tubercles, and ridges, and sensory tricorns, pit organs, pores, papillae and setae. Microscales are common features of the integument surface; their shape and size are highly variable. Tricorns were not observed on the marine littoral isopods. Several hitherto undescribed structures were observed including spade-like projections from the tergite surface of Oniscus asellus, hair-like filaments associated with the microscales of Jaera and ridged conical protuberances on Edotea triloba and E. montosa. The possible function of certain surface microstructures is discussed.     

Central projections from contact chemoreceptors of the locust ovipositor and adjacent cuticle

Contact chemoreceptors (basiconic sensilla) located on the ovipositor and genital segments of the locust serve to control the chemical features of the substrate before and during oviposition. They occur dispersed and also crowded in fields between mechanosensory exteroceptors sensitive to touch or wind (trichoid and filiform sensilla). The central nervous projections of the four chemosensory and one mechanosensory neurons from single basiconic sensilla were stained selectively, focusing on receptors on the ovipositor valves, which usually contact the substrate during the pre-oviposition probing movements. All axons and neurites from one contact chemoreceptor usually stay close together in most of their projections. Segregation occurs mainly when single axons terminate in one neuromere while the others proceed to a different neuromere or ganglion. For projections from one chemoreceptor, there is evidence neither for functional segregation of mechanosensory from chemosensory afferent terminals nor for specific segregation between different chemosensory afferents. The projections from sensilla of dorsal cuticle tend to project rather uniformly along the midline of the terminal ganglion. Comparative staining of touch- and wind-sensitive hair receptor neurons shows mostly central projections, similar to those of neighbouring contact chemoreceptors. From the typical intersegmental projections of most primary afferents and from the lack of segregation into glomerular structures, we conclude that integration of chemosensory information from the genital segments is distributed in the terminal and the 7th abdominal ganglion.     

Cuticular structures on the antennae of Hypoderma bovis De Geer (Diptera : Oestridae) females

The antennae of Hypoderma bovis (Diptera: Oestridae) females were examined using scanning electron microscopy. Each antenna is composed of 3 parts: the scape, the pedicel, and the funiculus, with a large, protruding arista. Mechanoreceptors are found on the proximal and lateral margins of the scape and pedicel, respectively. Microtrichia, which are presumably non-innervated, are located evenly over both the outer surface of the scape and the inner surface of the pedicel. A narrow band of microtrichia is present proximally on the outer surface of the pedicel. The entire funicular surface is densely covered with microtrichia. Small patches lacking these microtrichia appear as depressions or “pits” (8–20 μm in diameter) on the surface of the funiculus. Olfactory sensilla found on the funicular surface include basiconica type 1, basiconica type 2, and trichoid sensilla. The sensilla basiconica commonly occur in pits on the anterodorsal surface of the funiculus. Trichoid sensilla are abundant on the posteroventral surface of the funiculus and do not appear to occur in pits. In addition, clavate and peg sensilla, whose functions are unknown, are found in low numbers on the funicular surface. There may be as many as 300 olfactory pits on the anterodorsal surface of each funiculus. These are single-chambered and contain 6 or fewer sensilla basiconica. We propose that a relatively high number of pits may be characteristic of flies in Superfamily Oestroidea (as compared with those of Superfamily Muscoidea), but that pit morphology within the Calyptratae is not Superfamily-specific.