Sensory organs inIps typographus (Insecta: Coleoptera) — Fine structure of antennal sensilla

Summary The antennal sensilla inI. typographus are almost exclusively confined to the flattened terminal flagellar segment. The sensillar types have distinct distribution patterns in the three areas where they are found. Judging from the ultrastructural characteristics the following functions can be assigned to the sensillar types: chemoreception, single-walled and double-walled sensilla; chemoreception/mechanoreception, terminal-pore sensillum. Moreover there are two types of mechanoreceptors, one of which is connected to a bristle, whereas the other terminates within the cuticle of the flagellar segment.This study was made within the Swedish project Odour Signals for Control of Pest Insects.     

The sense of smell in Odonata: An electrophysiological screening

Volatile chemicals mediate a great range of intra- and interspecific signalling and information in insects. Olfaction has been widely investigated mostly in Neoptera while the knowledge of this sense in most basal insects such as Paleoptera (Odonata and Ephemeroptera) is still poor. In the present study we show the results of an electrophysiological screening on two model species, Libellula depressa (Libellulidae) and Ischnura elegans (Coenagrionidae), representatives of the two Odonata suborders Anisoptera and Zygoptera, with the aim to deep the knowledge on the sense of smell of this insect order. The antennal olfactory sensory neurons (OSNs) of these two species responded to the same 22 compounds (out of 48 chemicals belonging to different functional groups) encompassing mostly amines, carboxylic acids or aldehydes and belonging to green leaf volatiles, vertebrate related volatiles and volatiles emitted by standing waters bacteria. The properties of Odonata OSNs are very similar to those of ionotropic receptors (IRs) expressing OSNs in other insects.     

Electrophysiological identification of thermo- and hygro-sensitive receptor neurons on the antennae of the dragonfly Libellula depressa

Recent ultrastructural investigations on Odonata antennal flagellum describe two types of sensilla styloconica, T1 and T2. The styloconic sensilla are located in pits, at the bottom of deep cavities, and share common features typical of thermo-hygroreceptors. In order to ascertain if the Odonata antennae are involved in hygroreception and thermoreception, we carried out electrophysiological recordings (single cell recordings, SCR) from adult males and females of Libellula depressa L., 1758. After contact was established, the antenna was stimulated by rapid changes in temperature and humidity. The present research shows the occurrence of a dry (DC), a moist (MC) and a cold (CC) receptor neurons on the antennal flagellum of L. depressa. These data demonstrate for the first time the presence of functional thermo-hygroreceptors on the antennal flagellum of dragonflies. The present results extend our knowledge of the not visual sensory modalities of Odonata, a field of research unexplored so far.     

Hygroreceptors in the larva of Libellula depressa (Odonata: Libellulidae)

Ultrastructural and electrophysiological (single-cell recordings) investigations were carried out on the coeloconic sensilla borne by the apical antenna of the larvae of Libellula depressa (Odonata: Libellulidae). These sensilla appear as pegs located in pits. One of them is a compound sensillum constituted of two fused pegs in a common pit and the other two are single pegs located in separated pits close to each other. Coeloconic sensilla show position and ultrastructural details very similar to those described in insect hygroreceptors. The electrophysiological recordings on the apical antennae of the last larval instar of L. depressa clearly show the presence of moist and dry cells responding antagonistically to humidity changes. This study gives the first evidence of hygroreceptors in dragonfly larvae and represents the first electrophysiological approach to larval sensilla of aquatic insects. The presence of hygroreceptors in L. depressa larvae is in agreement with the hygropositive response shown by these insects in laboratory and field behavioural experiments.     

Volatile cues can drive the oviposition behavior in Odonata

Selection for the oviposition site represents the criterion for the behavioral process of habitat selection for the next generation. It is well known that in Odonata the most general cues are detected visually, but laboratory investigations on the coenagrionid Ischnura elegans showed through behavioral and electrophysiological assays that adults were attracted by olfactory cues emitted by prey and that males of the same species are attracted by female odor.The results of the present behavioral and electrophysiological investigations on I. elegans suggest the involvement of antennal olfactory sensilla in oviposition behavior. In particular, I. elegans females laid in the laboratory significantly more eggs in water from larval rearing aquaria than in distilled or tap water. Moreover, the lack of preference between rearing water and tap water with plankton suggests a role of volatiles related to conspecific and plankton presence in the oviposition site choice. I. elegans may rely on food odor for oviposition site selection, thus supporting the predictions of the “mother knows best” theory. These behavioral data are partially supported by electroantennographic responses. These findings confirm a possible role of olfaction in crucial aspects of Odonata biology.     

Asymmetric intraspecific competition among larvae of the damselfly Ischnura elegans (Zygoptera: Coenagrionidae)

Abstract. 1. A laboratory competition experiment is described in which the growth and development rates of larvae of the damselfly Ischnura elegans (Lind.) were measured over an entire instar.
2. Two larval instars which commonly occur together in the field were used in the experiment; they were maintained with a superabundance of prey and either larvae from the same or the larger/smaller instar.
3. Small larvae suffered increased development times and decreased size increases at the moult in the presence of large larvae but similar interference effects were not evident when these smaller larvae were in the presence of other small larvae.
4. Development time and size increases of large larvae were not significantly affected by the presence of small larvae.
5. Irrespective of the instar combinations investigated, interference effects were reduced when there were more perches available, although in only a few cases was this reduction significant.
6. The consequences of the asymmetric competition reported in the experiment for the study of lifetime reproductive success in damselflies are discussed. Late emerging adults may incur reduced reproductive success.     

The ecological role of caudal lamellae loss in the larval damselfly,Ischnura posita (Hagen) (Odonata: Zygoptera)

Summary Damselfly larvae may autotomize and regenerate any of their 3 caudal lamellae. At least one missing or regenerating lamella was evident in 50.1% of field collected Ischnura posita larvae. Lamellae loss during molting is very infrequent (1 out of 117 recorded molts). Laboratory trials indicate that conspecifics remove lamellae and that this process is density dependent. The percentage of larvae losing lamellae during 24 h trials ranged from 73.5 at the highest density tested to 17.3 at the lowest density. I. posita larvae are cannibalistic. The presence of lamellae reduces an individual's chance of being cannibalized. More than twice as many final instar lamellae-less larvae were cannibalized during 24 h trials than analogous individuals having 3 lamellae at experimental initiation. Costs are also associated with lamellae autotomy. 1) Although individuals without lamellae can swim they are more reluctant to release from a wooden stalk and swim when threatened (9% release) than are larvae with lamellae (29% release). Since swimming is part of their repertoire of anti-predator behaviors this behavioral shift should be detrimental. 2) Caudal lamellae function in O₂ uptake. Trials were conducted with larvae having and not having lamellae in an experimental horizontal oxygen gradient system. Relative to larvae without lamellae, those with lamellae preferred deeper depths at PO₂ values greater than 70 torr. Many lamellae-less larvae distributed themselves at the water surface throughout the range of PO₂ values tested. Differential depth distribution between larvae with and without lamellae is highly significant (P < 0.01).     

The hair-peg organs of the shore crab,Carcinus maenas (Crustacea,Decapoda): Ultrastructure and functional properties of sensilla sensitive to changes in seawater concentration

Summary The hair-peg organs of the shore crab, Carcinus maenas, are modified hair-sensilla. A small hair shaft (peg) is surrounded by a tuft of solid cuticular bristles (hairs). Each hair-peg organ is innervated by 6 sensory neurons, 2 of which have scolopidial (type-I) dendrites. The outer segments of all dendrites pass through a cuticular canal extending to the articulated hair base in which the 2 type-I dendrites terminate. The other 4 (type-II) dendrites reach the clavate tip of the hair shaft and have access to a terminal pore and a large sickle-shaped aperture. Three inner and 8–12 outer enveloping cells belong to a hair-peg organ. The innermost enveloping cell contains a scolopale, which has desmosomal connections to the ciliary rootlets of the type-I dendrites. An inner and an outer sensillum lymph space are present. The ultrastructural features of the dendrites and the cuticular apparatus indicate that the hair-peg organs are bimodal sensilla, comprising 2 mechano- and 4 chemosensitive sensory neurons. Extracellular recordings from the leg nerve indicate that the chemosensitive neurons of the hair-peg organs respond to changes in seawater concentration in the physiological range of Carcinus maenas.Supported by the Deutsche Forschungsgemeinschaft (SFB 45/A1; W. Gnatzy)     

Morphology and ultrastructure of the antennal chemoreceptors and mechanoreceptors of worker Coptotermes formosanus Shiraki

Summary Three types of setae were found on the antenna of workers of C. formosanus. Sensilla basiconica function as chemoreceptors. They are non-socketed, with fixed plate base, thin walled, and perforated. Pore tubules are contained within the cuticular pores of s. basiconica. The pore tubules have direct contact with the dendritic branches within the sensilla. The other sensilla are tactile mechanoreceptors. The longest setae (sensilla trichodea) are thick walled, socketed, mononeuronic, and non-perforated. The shorter mechanoreceptors (sensilla chaetica) have the same characteristics as s. trichodea, exept that they are sometimes dineuronic. The numbers and distribution of the sensilla were established.Supported by the College of Agricultural and Life Sciences, University of Wisconsin, Madison, and in part by the M.U.C.I.A. program, the National Science Foundation Grant No. 36700, and the U.S.D.A., Forest Service Cooperative Project 12-13. The authors also acknowledge the SEM and TEM assistance of S. D. Carlson, M. Garment, and G. Gaard, respectively     

The effects of temperature and prey density on the development rates and growth of damselfly larvae (Odonata: Zygoptera)

Abstract. 1. The effects of prey density and temperature on the feeding and development rates of several late instars of the larvae of three common European damselflies ( Lestes sponsa (Hansemann), Coenagrion puella (L.) and Ischnura elegans van der Linden) were investigated in a laboratory experiment.
2. Functional responses were used to estimate maximum feeding rates. Maximum feeding rates were compared between species, instars and temperatures by expressing prey consumption in terms of prey biomass consumed per unit predator biomass. Lestes was capable of feeding at almost twice the rate of either Coenagrion or Ischnura.
3. Higher feeding rates led to faster development rates and there was an interaction between species and temperature. With the exception of those with very low feeding rates, larvae maintained at higher temperatures, but similar feeding rates, developed faster. Under similar conditions of temperature and feeding rate, Lestes larvae developed faster than larvae of either Coenagrion or Ischnura.
4. Faster development rates at similar rates of food intake were achieved at the cost of reduced size-increases between instars.
5. The differences in the responses of the three species are discussed in the light of their respective life histories, and with reference to a recent model of population regulation in damselflies.     

The eversible tentacle organs of Polyommatus caterpillars (Lepidoptera,Lycaenidae): Morphology,fine structure,sensory supply and functional aspects

In their late (3rd and 4th) larval stages, caterpillars of the myrmecophilous lycaenid (Lepidoptera) species Polyommatus coridon and Polyommatus icarus, possess on their 8th abdominal segment two eversible so called tentacle organs (TOs). Previous histological and behavioural results have proposed that the TOs may release a volatile substance that elicits “excited runs” in attendant ants. In our study we investigated for the first time the temporal in- and eversion pattern of TOs. Using nerve tracing, Micro-CT, light- and electron microscopy techniques we studied (i) the histology of the 8th abdominal segment, (ii) the fine structure of the cuticular and cellular apparatus of the TOs, (iii) the attachment sites of the retractor muscle of each TO and (iv) the fine structure of the long slender tentacle hairs which are exposed to the outside, when the TOs are everted and fold back into the TO-sac during inversion. Our data show that the tentacle hairs are typical insect mechanoreceptors, each innervated by a small bipolar sensory cell with a tubular body in the tip of the outer dendritic segment. The latter is enclosed by a cuticular sheath previously called the “internal cuticular duct” and misinterpreted in earlier studies as the space, where the tentacle hairs actively secrete fluids. However, we found no glandular structures nearby or in the wall of the TO-sac. Also we did not reveal any conspicuous signs of secretory activity in one of the enveloping cells belonging to a tentacle hair. Although highly unusual features for an insect mechanoreceptor are: (a) the hair-shaft lumen of tentacle hairs contains flocculent material as well small vesicles and (b) the thin cuticular wall of the hair-shaft and its spines possess few tiny pores. Our data do not support the assumption of previous studies that volatile substances are released via the tentacle organs during their interactions with ants which in turn are supposed to cause excited runs in ants.     

Lamellated outer dendritic segments of a chemoreceptor within wall-pore sensilla in the labial palp-pit organ of the butterfly,Pieris rapae L. (Insecta,Lepidoptera)

Summary The structure of the sensilla in the apical pit of the third segment of the labial palps in Pieris rapae was investigated in cryofixed and chemically fixed specimens. There is a field of about 80 club-shaped sensilla, 94% of which house a single sensory cell; 6% contain two sensory cells. All sensory cells are of the same type and are characterized by the structure of the dendritic outer segment. This consists of a proximal cylindrical and a distal lamellated section. The lamellae contain a lattice of longitudinally arranged microtubules. Filamentous strands connect the microtubules with the surface membrane of the lamellae. The surface area of the lamellated section amounts to about 40 m². Pores and pore tubules are present in the cuticular wall of the peg. Electrophysiological recordings show that the sensory cells are olfactory receptors, which react to a variety of complex plant odors and to the odor of conspecifics. It is shown that (a) the usual modality-specific characteristics of insect olfactory sensilla apply here also; (b) lamellation is not only a characteristic of thermoreceptors, but also of olfactory chemoreceptors; (c) there are pore tubules that are separated from the dendritic membranes by an extended dendritic sheath; and (d) in the labial palppit sensilla only the lamellated dendritic tip region may be involved in sensory transduction.Supported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

Are the funnel-canal organs the ‘campaniform sensilla’ of the shore crab,Carcinus maenas (Decapoda,Crustacea)?

Summary The funnel-canal organs on the dactyls of the shore crab, Carcinus maenas, are innervated by 3–24 sensory cells with unbranched dendrites, which attain a length of 500–1400 m. The outer dendritic segments are enclosed in a dendritic sheath and pass through the cuticle within a canal. Two dendrite types can be distinguished according to ultrastructural criteria: Type I has a long ciliary segment, A-tubules with an osmiophilic core and arms, and a thick ciliary rootlet. Type II possesses only a short ciliary segment and a thin ciliary rootlet. Each funnel-canal organ contains two type-I dendrites. Their ciliary bases appear a few m distal to those of the type-II dendrites (1 to 22 in number). Two inner and two to eight outer enveloping cells belong to a sensillum. The innermost enveloping cell contains a large scolopale. In the second enveloping cell single scolopale rods are present. Thus, the funnel-canal organs are characterized by structural features typical for mechano-sensitive scolopidia, on the one hand, and for chemoreceptors, on the other. Therefore, the funnel-canal organs are very likely bimodal sensilla (contact chemoreceptors). A comparison with other arthropod sensilla shows that cuticular mechanoreceptors of aquatic crustaceans generally exhibit a scolopidial organization.     

Survival cost of an intrasexually selected ornament in a damselfly

Ornaments could evolve as honest indicators of fighting ability, provided they have costs that make deceptive signalling unprofitable. I tested for such costs by manipulating the size of the intrasexually selected wing spots of male rubyspot damselflies (Hetaerina americana) and monitoring survival in the field. Males with enlarged spots had higher mortality rates than both unmanipulated and sham-manipulated controls. Natural wing spot size correlated positively with longevity, which suggests that higher quality males develop larger spots.     

Ultrastructure of the chemosensitive basiconic single-walled wall-pore sensilla on the antennae in adults and embryonic stages of Locusta migratoria L. (Insecta,Orthoptera)

Summary The structure and embryonic development of the two types (A, B) of basiconic sensilla on the antennae of Locusta migratoria were studied in material that had been cryofixed and freeze-substituted, or chemically fixed and dehydrated. Both types are single-walled wall-pore sensilla. Type-A sensilla comprise 20–30 sensory and 7 enveloping cells. One enveloping cell (thecogen cell secretes the dendrite sheath); four are trichogen cells, projections of which form the trichogen process during the 2nd embryonic molt. The trichogen cells form two concentric pairs proximally. Two tormogen cells secrete the cuticular socket of the sensillum. The dendritic outer segments of the sensory cells are branched. Bifurcate type-A sensilla have also been observed. Type-B sensilla comprise three sensory and four enveloping cells (one thecogen, two trichogen and one tormogen). The trichogen process is formed by the two trichogen cells, each of which gives rise to two projections. The trichogen cells are concentrically arranged. The dendritic outer segments of the sensory cells are unbranched. In the fully developed sensillum, all trichogen and tormogen cells border on the outer receptor lymph cavity. It is suggested that the multicellular organization of the type-A sensilla can be regarded as being advanced rather than primitive.Supported by the Dcutschc Forschungsgemeinschaft (SFB 4/G1)     

Mechanical and tactile incompatibilities cause reproductive isolation between two young damselfly species

External male reproductive structures have received considerable attention as a cause of reproductive isolation (RI), because the morphology of these structures often evolves rapidly between populations. This rapid evolution presents the potential for mechanical incompatibilities with heterospecific female structures during mating and could thus prevent interbreeding between nascent species. Although such mechanical incompatibilities have received little empirical support as a common cause of RI, the potential for mismatch of reproductive structures to cause RI due to incompatible species‐specific tactile cues has not been tested. We tested the importance of mechanical and tactile incompatibilities in RI between Enallagma anna and E. carunculatum, two damselfly species that diverged within the past ～250,000 years and currently hybridize in a sympatric region. We quantified 19 prezygotic and postzygotic RI barriers using both naturally occurring and laboratory‐reared damselflies. We found incomplete mechanical isolation between the two pure species and between hybrid males and pure species females. Interestingly, in mating pairs for which mechanical isolation was incomplete, females showed greater resistance and refusal to mate with hybrid or heterospecific males compared to conspecific males. This observation suggests that tactile incompatibilities involving male reproductive structures can influence female mating decisions and form a strong barrier to gene flow in early stages of speciation.     

Environmental heterogeneity and alternative mating tactics in the damselfly Protoneura amatoria

Conditional male mating strategies have been studied extensivelyin relation to male attributes, such as size and resource-holdingpotential, but few studies have considered the effects of environmentalheterogeneity on the use of alternative mating tactics. In somesystems, environmental heterogeneity may be the key to understandingthe evolution and maintenance of such polyphenisms. I examinedthe influence of the physical environment on the use of alternativetactics by the damselfly Protoneura amatoria. Male P. amatoriareversibly use 2 tactics to gain matings: 1) sit and wait inthe canopy for passing females or 2) hover over the water andattempt to grab females that are ovipositing in floating debris.Observations in 3 streams indicated that the use of the hoveringtactic was greater under high-light than low-light conditionsand at higher densities of ovipositing females. The densityof ovipositing females was correlated with both the light conditionsand the availability of oviposition substrate, indicating thatphysical factors exert indirect as well as direct influenceson tactic use. Experimental manipulations showed that both malesand females responded directly to light conditions and suggestedthat males responded directly to the density of ovipositingfemales. These results can be explained largely in terms ofthe cues and constraints inherent in different light environments.Thus, the conditional mating strategy of P. amatoria appearsto have evolved in response to, and been maintained by, fine-scalevariation in the physical environment. These findings are discussedin relation to flight dynamics and predation risk.     

Size-biased dispersal prior to breeding in a damselfly

Summary Dispersal is notoriously difficult to measure, so its potential population consequences are often unknown. If dispersal is density-dependent, it can act in population regulation. Adult damselflies Enallagma boreale (Zygoptera: Coenagrionidae) raised as larvae under a range of competitive regimes were individually measured and marked. Individuals that survived to reproductive maturity were either recovered at the natal pond or had dispersed to nearby water bodies. Dispersing individuals were heavier at emergence than those returning to the natal pond to breed. Therefore, an increased probability of dispersal does not appear to be a response to poor conditions in this species.     

Pedestal hairs of the ant Echinopla melanarctos (Hymenoptera, Formicidae): morphology and functional aspects

The South East Asian arboreal Formicine Echinopla melanarctos, as well as some other members of this genus possess a cuticular structure unique in ants, the pedestal hairs. In E. melanarctos, about 700 pedestal hairs are situated on the dorsal and lateral surfaces of the head, the alitrunk, the petiole and the gaster. They are arranged in a polygon-like figuration. On the summit of each of the up to 200-μm high pedestals, a single central hair inserts. This hair (up to 500-μm long) is innervated by a single bipolar mechanosensitive sensory cell. The lumen of each tube-like pedestal contains (1) epithelial cells (2) the sensory cell and the auxiliary cells of the central hair and (3) the long efferent ductules of up to ten isolated bicellular glandular units. Each glandular unit is composed of a secretory glandular cell and a duct cell, all of which are located at the base of a pedestal. The cytoplasm of a glandular cell contains a well-developed end apparatus and is characterised by stacks of smooth and granular endoplasmic reticulum, numerous polyribosomes, a lot of mitochondria and some up to 5-μm large secretory vesicles. The secretion of the gland cells is released on the apex of the pedestal wall via small pores. Approximately 30 μm below their summit, some pedestals possess additionally (up to six) mechanosensitive hairs that are arranged ray-like. We suppose that the pedestal hairs are important in nest-space protection and find that only in ants with high pedestals on the head (Echinopla melanarctos and Echinopla pallipes), the compound eyes are stalked thus overtopping the pedestals.     

Sexual selection and temporal phenotypic variation in a damselfly population

Temporal variation in selection can be generated by temporal variation in either the fitness surface or phenotypic distributions around a static fitness surface, or both concurrently. Here, we use within- and between-generation sampling of fitness surfaces and phenotypic distributions over 2 years to investigate the causes of temporal variation in the form of sexual selection on body size in the damselfly Enallagma aspersum. Within a year, when the average female body size differed substantially from the average male body size, male body size experienced directional selection. In contrast, when male and female size distributions overlapped, male body size experienced stabilizing selection when variances in body size were large, but no appreciable selection when the variances in body size were small. The causes of temporal variation in the form of selection can only be inferred by accounting for changes in both the fitness surface and changes in the distribution of phenotypes.