Functional Diversity of Vibrational Signaling Systems in Insects.

Communication by substrate-borne mechanical waves is widespread in insects. The specifics of vibrational communication are related to heterogeneous natural substrates that strongly influence signal transmission. Insects generate vibrational signals primarily by tremulation, drumming, stridulation, and tymbalation, most commonly during sexual behavior but also in agonistic, social, and mutualistic as well as defense interactions and as part of foraging strategies. Vibrational signals are often part of multimodal communication. Sensilla and organs detecting substrate vibration show great diversity and primarily occur in insect legs to optimize sensitivity and directionality. In the natural environment, signals from heterospecifics, as well as social and enemy interactions within vibrational communication networks, influence signaling and behavioral strategies. The exploitation of substrate-borne vibrational signaling offers a promising application for behavioral manipulation in pest control.

Stink Bug Communication and Signal Detection in a Plant Environment.

Plants influenced the evolution of plant-dwelling stink bugs' systems underlying communication with chemical and substrate-borne vibratory signals. Plant volatiles provides cues that increase attractiveness or interfere with the probability of finding a mate in the field. Mechanical properties of herbaceous hosts and associated plants alter the frequency, amplitude, and temporal characteristics of stink bug species and sex-specific vibratory signals. The specificity of pheromone odor tuning has evolved through highly specific odorant receptors located within the receptor membrane. The narrow-band low-frequency characteristics of the signals produced by abdomen vibration and the frequency tuning of the highly sensitive subgenual organ vibration receptors match with filtering properties of the plants enabling optimized communication. A range of less sensitive mechanoreceptors, tuned to lower vibration frequencies, detect signals produced by other mechanisms used at less species-specific levels of communication in a plant environment. Whereas the encoding of frequency-intensity and temporal parameters of stink bug vibratory signals is relatively well investigated at low levels of processing in the ventral nerve cord, processing of this information and its integration with other modalities at higher neuronal levels still needs research attention.

Female competition for availability of males in insects: the Nezara viridula (Linnaeus, 1758) model.

Multimodal communication in solitary stinkbugs enables them to meet, mate and copulate. Many plant-dwelling species exchange information during the calling phase of mating behavior using substrate-borne vibratory signals. A female-biased gender ratio induces rivalry and competition for a sexual partner. Female competition for males, first described among Heteroptera in three stinkbug species, revealed species specific differences and opened the question of plasticity in individually emitted temporal and frequency signal characteristics during calling and rival alternation. To address this question and gain an insight into the mechanisms underlying stinkbug female rivalry, we compared the characteristics of alternated signals in the southern green stinkbug Nezara viridula (Linnaeus, 1758) (Hemiptera: Pentatomidae). Compared to male rivalry, female rivalry is more complex, lasts longer and runs through successive phases by a combination of different song types. The male pheromone triggers alternation between females, producing song pulses that occasionally overlap each other. One female initiates the rivalry by changing individual pulses into pulse trains of three different types. The competing female alternates with pulses of changed temporal characteristics at lower levels of rivalry and by varying the frequency characteristics of pulse trains at higher levels. During female rivalry, the male either stops responding or occasionally emits calling and courtship signals in response to the female that has produced signals of steady temporal characteristics. Female rivalry shows complex and species specific patterns of information exchange at different levels with a broad-range variation of temporal and frequency characteristics of, until now, unidentified vibratory emissions.

A male-produced aggregation-sex pheromone of the beetle Arhopalus rusticus (Coleoptera: Cerambycidae, Spondylinae) may be useful in managing this invasive species.

The longhorned beetle Arhopalus rusticus (Coleoptera: Cerambycidae, Spondylinae) is a common species in conifer forests of the Northern Hemisphere, but with global trade, it has invaded and become established in New Zealand, Australia, and South America. Arhopalus rusticus is a suspected vector of the phytopathogenic nematode, Bursaphelenchus xylophilus, the causative agent of pine wilt disease, which is a major threat to pine forests worldwide. Here, we report the identification of a volatile, male-produced aggregation-sex pheromone for this species. Headspace odours from males contained a major male-specific compound, identified as (2 S, 5E)-6,10-dimethyl-5,9-undecadien-2-ol (common name (S)-fuscumol), and a minor component (E)-6,10-dimethyl-5,9-undecadien-2-one (geranylacetone). Both compounds are known pheromone components for species in the same subfamily. In field trials in its native range in Slovenia, (S)-fuscumol was significantly more attractive to beetles of both sexes, than racemic fuscumol and a blend of host plant volatiles commonly used as an attractant for this species. Fuscumol-baited traps also caught significant numbers of another spondylidine species, Spondylis buprestoides (L.), and a rare click beetle, Stenagostus rufus (De Geer). The pheromone can be exploited as a cost-effective and environmentally safe tool for detection and monitoring of this invasive species at ports of entry, and for monitoring the beetle's distribution and population trends in both endemic and invasive populations.

Olfactory signaling of aggressive intent in male-male contests of cave crickets (Troglophilus neglectus; Orthoptera: Rhaphidophoridae).

In animal contests, communicating aggressive motivation is most often mediated by visual or acoustic signals, while chemical signals are not expected to serve such a function since they are less able to be modulated by the sender during the changing behavioral context. We describe a rare example of ephemeral olfactory signals in terrestrial animals, signals that are emitted via protrusive scent glands in male cave crickets Troglophilus neglectus (Orthoptera, Rhaphidophoridae) to reflect the state of the signaler's aggression. We correlate the intensity of behaviorally expressed aggression of the individuals in dyadic contests with the frequency and extent of their gland tissue protrusion, the latter serving as an indication of the amount of released odor. We detected large amounts of odor release during brief gland protrusions, and the absence of its release during gland retraction. Males protruded the glands during and after encountering a rival, with the degree of protrusion increasing with the intensity of the signalers' aggression. During the encounters, the degree of gland protrusion increased most strongly with the occurrence of the elevated body posture, directly preceding the attack. This degree was significantly higher in encounter winners than in losers displaying such posture, suggesting the highly important role of the released odor for contest resolution. After the encounters, glands were protruded almost exclusively by winners, apparently announcing victory. We tested for the function of the olfactory signals also directly, by preventing gland tissue protrusion in symmetric and asymmetric treatments of the contestants. Treating only the dominant individuals decreased the percentage of encounters they won by over 60%, while treating both contestants elicited a significant increase in the frequency and duration of fights. During contests, the olfactory signals of T. neglectus apparently function as a highly effective threat, which prevents maximal contest escalation and decreases the conflict-related costs.

Novel, male-produced aggregation pheromone of the cerambycid beetle Rosalia alpina, a priority species of European conservation concern.

Several recent studies have demonstrated the great potential for exploiting semiochemicals in ecology and conservation studies. The cerambycid beetle Rosalia alpina represents one of the flagship species of saproxylic insect biodiversity in Europe. In recent years its populations appear to have declined substantially, and its range has shrunk considerably as a result of forest management and urbanization. Here, we collected volatile chemicals released by males and females of R. alpina. Analyses of the resulting extracts revealed the presence of a single male-specific compound, identified as a novel alkylated pyrone structure. In field bioassays in Slovenia, traps baited with the synthesized pyrone captured both sexes of R. alpina, indicating that the pyrone functions as an aggregation pheromone. Our results represent the first example of a new structural class of pheromones within the Cerambycidae, and demonstrate that pheromone-baited traps can provide a useful tool for sampling R. alpina. This tool could be particularly useful in the ongoing development of conservation strategies for the iconic but endangered Alpine longicorn.

Male adaptations to minimize sexual cannibalism during reproduction in the funnel-web spider Hololena curta.

Males of many spider species risk being attacked and cannibalized while searching for, courting, and mating with conspecific females. However, there are exceptions. We show that the funnel-web spider, Hololena curta, has 3 adaptations that minimize risk to males during courtship and mating, and enhance reproductive success. First, males detected chemical or tactile signals associated with webs of virgin females, and differentiated them from webs of mated females, enabling males to increase encounter rates with virgin females and avoid aggressive mated females. Second, males produced stereotyped vibrational signals during courting which induced female quiescence and suppressed female aggression. Third, when touched by males, sexually receptive females entered a cataleptic state, allowing males to safely approach and copulate. Because males can mate multiple times and the sex ratio in natural populations of H. curta is female biased, overall reproductive output is likely increased by males of this species avoiding sexual cannibalism.

Interference of Overlapping Insect Vibratory Communication Signals: An Eushistus heros Model.

Plants limit the range of insect substrate-borne vibratory communication by their architecture and mechanical properties that change transmitted signal time, amplitude and frequency characteristics. Stinkbugs gain higher signal-to-noise ratio and increase communication distance by emitting narrowband low frequency vibratory signals that are tuned with transmission properties of plants. The objective of the present study was to investigate hitherto overlooked consequences of duetting with mutually overlapped narrowband vibratory signals. The overlapped vibrations of the model stinkbug species Eushistus heros, produced naturally or induced artificially on different plants, have been analysed. They represent female and male strategies to preserve information within a complex masked signal. The brown stinkbugs E. heros communicate with species and gender specific vibratory signals that constitute characteristic duets in the calling, courtship and rivalry phases of mating behaviour. The calling female pulse overlaps the male vibratory response when the latency of the latter is shorter than the duration of the female triggering signal or when the male response does not inhibit the following female pulse. Overlapping of signals induces interference that changes their amplitude pattern to a sequence of regularly repeated pulses in which their duration and the difference between frequencies of overlapped vibrations are related inversely. Interference does not occur in overlapped narrow band female calling pulses and broadband male courtship pulse trains. In a duet with overlapped signals females and males change time parameters and increase the frequency difference between signals by changing the frequency level and frequency modulation pattern of their calls.