Stridulation und Kommunikation durch Substratschall beiGecarcinus lateralis (Crustacea Decapoda)

Zusammenfassung Ein für echte Landkrabben (Familie Gecarcinidae) bisher unbekanntes Kommunikationssystem wird fürGecarcinus lateralis beschrieben. Die Informationsinhalte Drohen, Besänftigen und Balzen werden durch regelmäßig aufeinanderfolgende Substratschallimpulse übermittelt, wobei der jeweilige Informationsinhalt durch die Anzahl der Impulse pro Zeit gekennzeichnet ist. Die Erzeugung von Substratschall durch Stridulation ist Bestandteil dieses Kommunikationssystems.

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Stridulation and communication by substrate vibration inGecarcinus lateralis (Crustacea Decapoda)

Summary In the terrestrial crabGecarcinus lateralis (Gecarcinidae) a new type of communication system was found. Information is transmitted by substrate vibration. Typical sequences of pulses were emitted by one crab in order to inform the receiving crab about the senders threatening, appeasing or sexual display behavior. The number of pulses per time unit was different for each type of behavior pattern. Stridulation transmitted as substrate vibration is also part of this communication system.

     

Foraging vibration signals attract foragers and identify food size in the drywood termite, <Emphasis Type="Italic">Cryptotermes secundus</Emphasis>

Information gathering and communication behaviour has evolved within constraints of size, physiology and ecology of the animal. Due to these constraints, small herbivorous insects are likely to use substrate borne vibrations for information gathering and communication. Although such signals have been characterised in many types of insects, including group-living insects, they are poorly known in termites.We showed that the Australian drywood termite Cryptotermes secundus could determine the size of wooden blocks by using the vibrations generated during foraging. The termites behaved differently in choice experiments when artificially generated vibration signals were played compared with natural recordings, indicating that these termites can discriminate the source of the vibration as well. AT-maze experiment showed that the termites were attracted to the natural recordings of feeding termites, suggesting that vibrations are important in communication during foraging as well as food resource assessment. Combining the effects of food size preference and attraction to other termites explained differences in behaviour between artificially generated vibration signals compared with natural recordings. This study demonstrates that termites use substrate borne vibrations for information gathering and communication as predicted. Received 21 March 2007; revised 19 June and 7 August 2007; accepted 20 August 2007.     

The signaller's dilemma: a cost-benefit analysis of public and private communication

Background

Understanding the diversity of animal signals requires knowledge of factors which may influence the different stages of communication, from the production of a signal by the sender up to the detection, identification and final decision-making in the receiver. Yet, many studies on signalling systems focus exclusively on the sender, and often ignore the receiver side and the ecological conditions under which signals evolve.

Methodology/Principal Findings

We study a neotropical katydid which uses airborne sound for long distance communication, but also an alternative form of private signalling through substrate vibration. We quantified the strength of predation by bats which eavesdrop on the airborne sound signal, by analysing insect remains at roosts of a bat family. Males do not arbitrarily use one or the other channel for communication, but spend more time with private signalling under full moon conditions, when the nocturnal rainforest favours predation by visually hunting predators. Measurements of metabolic CO₂-production rate indicate that the energy necessary for signalling increases 3-fold in full moon nights when private signalling is favoured. The background noise level for the airborne sound channel can amount to 70 dB SPL, whereas it is low in the vibration channel in the low frequency range of the vibration signal. The active space of the airborne sound signal varies between 22 and 35 meters, contrasting with about 4 meters with the vibration signal transmitted on the insect''s favourite roost plant. Signal perception was studied using neurophysiological methods under outdoor conditions, which is more reliable for the private mode of communication.

Conclusions/Significance

Our results demonstrate the complex effects of ecological conditions, such as predation, nocturnal ambient light levels, and masking noise levels on the performance of receivers in detecting mating signals, and that the net advantage or disadvantage of a mode of communication strongly depends on these conditions.     

First Data on Vibration Signals in Flies of the Genus Meromyza (Diptera,Chloropidae)

Biology Bulletin - This paper discusses vibration communication in representatives of the genus Meromyza. The frequency range of vibration signals of Meromyza saltatrix (L., 1761) females is 229 to...     

Structure and sensory physiology of the leg scolopidial organs in Mantophasmatodea and their role in vibrational communication

Individuals of the insect order Mantophasmatodea use species-specific substrate vibration signals for mate recognition and location. In insects, substrate vibration is detected by mechanoreceptors in the legs, the scolopidial organs. In this study we give a first detailed overview of the structure, sensory sensitivity, and function of the leg scolopidial organs in two species of Mantophasmatodea and discuss their significance for vibrational communication. The structure and number of the organs are documented using light microscopy, SEM, and x-ray microtomography. Five scolopidial organs were found in each leg of male and female Mantophasmatodea: a femoral chordotonal organ, subgenual organ, tibial distal organ, tibio-tarsal scolopidial organ, and tarso-pretarsal scolopidial organ. The femoral chordotonal organ, consisting of two separate scoloparia, corresponds anatomically to the organ of a stonefly (Nemoura variegata) while the subgenual organ complex resembles the very sensitive organs of the cockroach Periplatena americana (Blattodea). Extracellular recordings from the leg nerve revealed that the leg scolopidial organs of Mantophasmatodea are very sensitive vibration receptors, especially for low-frequency vibrations. The dominant frequencies of the vibratory communication signals of Mantophasmatodea, acquired from an individual drumming on eight different substrates, fall in the frequency range where the scolopidial organs are most sensitive.     

Intra-Patriline Variability in the Performance of the Vibration Signal and Waggle Dance in the Honey Bee, Apis mellifera

We examined intra-patriline behavioral plasticity in communication behavior by generating lifetime behavioral profiles for the performance of the vibration signal and waggle dance in workers which were the progeny of three unrelated queens, each inseminated with the semen of a single, different drone. We found pronounced variability within each patriline for the tendency to produce each signal, the ontogeny of signal performance, and the persistence with which individual workers performed the signals throughout their lifetimes. Within each patriline, the number of workers that performed each signal and the distribution of onset ages for each signal were significantly different. In each patriline, workers of all ages could perform vibration signals; vibration signal production began 3–5 d before waggle dancing; and some workers began performing waggle dances at ages typically associated with precocious foraging. Most workers vibrated and waggled only 1–2 d during their lifetimes, although each patriline contained some workers that performed the signal persistently for up to 8 or 9 d. We also found marked variability in signal performance among the three worker lineages examined. Because the vibration signal and waggle dance influence task performance, variability in signaling behavior within and between subfamilies may help to organize information flow and collective labor in honey bee colonies. Inter-patriline variability may influence the total number of workers from different partrilines that perform the signals, whereas intra-patriline variability may further fine-tune signal performance and the allocation of labor to a given set of circumstances. Although intra-patriline behavioral variability is assumed to be widespread in the social insects, our study is the first to document the extent of this variability for honey bee communication signals.     

Substrate-Borne Vibration Mediates Intrasexual Agonism in the New Zealand Cook Strait Giant Weta (<Emphasis Type="Italic">Deinacrida rugosa</Emphasis>)

Substrate-borne vibrational communication is a common mode of information transfer in many invertebrate groups, with vibration serving as both primary and secondary signal channels in Orthopterans. The Cook Strait giant weta, Deinacrida rugosa (Orthoptera: Anostostomatidae), is an endangered New Zealand insect whose communication system has not been previously described. After field observations of intraspecific interactions in D. rugosa provided preliminary evidence for substrate-borne vibrational communication in the species, we sought to identify the following: vibrational signal structure, the mechanism of signal production, whether signal production is a sexually dimorphic trait, whether substrate-borne signals encode information regarding sender size, the primary social context in which vibration is utilized and finally, the function of vibrational signaling in the species. We used laser Doppler vibrometry to show that D. rugosa males produce low frequency (DF?=?37.00?±?1.63 Hz) substrate-borne vibrations through dorso-ventral tremulation. Rarely produced by females, male signals appear to target rivals while both are in the direct physical presence of a female. Tremulatory responses to playbacks were only produced by males in male-male-female trial contexts, and neither sex exhibited walking vibrotaxis to playback signals, indicating that substrate-borne vibrational signals are not likely a component of the courtship repertoire. While we found that vibrational signal structure was not closely related to signaler size, males that initiated male-male signaling bouts held a significant advantage in contests.     

Vibration and Animal Communication: A Review

Vibration through the substrate has likely been important toanimals as a channel of communication for millions of years,but our awareness of vibration as biologically relevant informationhas a history of only the last 30 yr. Morphologists know thatthe jaw mechanism of early amphibians allowed them to perceivevibration through the substrate as their large heads lay onthe ground. Although the exact mechanism of vibration productionand the precise nature of the wave produced are not always understood,recent technical advances have given answers to increasinglysophisticated questions about how animals send and receive signalsthrough the substrate. Some of us have been forced to explorethe use of vibration when all other attempts to manipulate animalsin the field have failed, while others began to think aboutvibration to explain some of the puzzling behaviors of speciesthey were studying in other contexts. It has thus become clearthat the use of vibration in animal communication is much morewidespread than previously thought. We now know that vibrationprovides information used in predator-prey interactions, recruitmentto food, mate choice, intrasexual competition and maternal/broodsocial interactions in a range of animals from insects to elephants.     

Age and Behavior of Honey Bees,Apis mellifera (Hymenoptera: Apidae), that Perform Vibration Signals on Workers

The vibration signal is one of the most commonly occurring communication displays in honey bee (Apis mellifera) colonies. It may function in a ‘modulatory’ manner, because it causes a nonspecific increase in activity that enhances a variety of behaviors depending upon the age and caste of the recipient. We examined honey bee workers that performed vibration signals on other workers in three observation hives, each containing a population of marked bees of known age. In all three colonies, the mean age of the first performance of the vibration signal was significantly different from the mean age at which workers first performed waggle dances, carried pollen loads, or attended the queen. However, workers of all ages, except those less than 3 d old, could perform vibration signals. In older workers of foraging age, signal performance was most closely associated with recent foraging success. Younger workers that vibrated did not appear to be early-maturing foragers and thus their signals were probably not influenced by food collection. Rather, for these preforaging-age workers, signal performance was associated more with periods of orientation flight, during which younger bees learn the location of the nest and surrounding landmarks. Thus, the vibration signal may be triggered by different stimuli in different worker age classes. Because it elicits a general increase in activity in all recipients, the signal may help adjust many different colony behaviors simultancously to changes in foraging success and colony development.     

Palomena prasina (Hemiptera: Pentatomidae) vibratory signals and their tuning with plant substrates

Palomena prasina is interesting for the study of vibrational communication within the Pentatomid subfamily Pentatominae, because its host range is limited to woody plants, unlike the better known Nezara viridula, whose vibrational communication is commonly used as a model for the whole family. The vibrational repertoire of P. prasina was described several decades ago and is redescribed in this paper using modern methods for non-contact vibration recording. Additionally, we hypothesized that this species has retained the capacity for signal frequency variation necessary for tuning to resonance properties of various host plants of Pentatominae, but if the signals are emited in the absence of mechanical feedback, they are tuned more specifically to their native acoustic environment — woody plants. By recording live bugs signalling on different substrates and comparing spectral properties of their signals among substrates, we found that there is a match between the signals emitted on a woody branch and those emitted on a non-resonant surface, while spectral properties of signals emitted on herbaceous plants differ. Our findings provide evidence in support of the signal tuning hypothesis and shed further light on the crucial role of substrate in vibrational communication of insects.     

The Role of the Vibration Signal during Nest‐Site Selection by Honey Bee Swarms

The vibration signal may influence nest‐site selection by honey bee swarms by enhancing scouting and recruitment. We investigated this hypothesis by comparing (1) the number of nest sites and the distances communicated by nest‐site dancers on swarms from which vibrators were and were not removed and (2) the behavior of scouts visiting higher‐quality (HQ) and lower‐quality (LQ) nest boxes. The removal of vibrators from swarms did not alter the number of nest sites investigated, the distances traveled to nest sites, or the time required to select a new nest cavity. In contrast, vibrator removal tripled the time required for swarms to achieve liftoff after a cavity had been selected, although all swarm eventually became airborne and moved to a new site. About 14% of the scouts that visited the HQ and LQ nest boxes performed vibration signals; however, nest‐box quality did not influence the tendency to produce the signal or intermix vibration signals and recruitment dances. However, we did find a significant, positive correlation between overall levels of vibration signal activity and nest‐site recruitment during the house‐hunting process. When viewed in concert, our results suggest that the vibration signal contributes to the house‐hunting process by operating in a non‐specific manner that may enhance scouting and recruitment in general during nest‐site selection and facilitate rapid swarm liftoff after a new nest site has been chosen. The vibration signal is therefore a component in the cascade of communication signals that orchestrate house‐hunting and colony relocation decisions.     

The sounds produced by flies and bees

Summary The sounds produced by the thoracic flight machinery of bees and flies appear to be composed of two main vibration modes. The lower frequency one corresponds to the wingbeat frequency. The higher frequency vibration, which is in the kcps. range, is reset in phase on every wingbeat half-cycle. Therefore Sonagraphic or Fourier analysis of the sounds gives only harmonics of the wingbeat frequency. However, oscillograms of the waveforms show that the higher frequency vibration is nearly independent of wingbeat frequency.The high frequency vibration is probably important in bee communication. We speculate that it is due to skeletal vibration which is relatively undamped by muscular and aerodynamic loading.

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Zusammenfassung Der Thorax von Insekten, deren Flugmuskeln einen myogenen Rhythmus zeigen, vibriert in Aktivitätsphasen meist mit zwei voneinander unabhängigen Frequenzen: Die erste wird durch die elastischen Eigenschaften des Skelets, die Kontraktionen der indirekten Flugmuskeln und die Belastung der Flügel bestimmt (Flügelschlagfrequenz). Sie kann über das Nervensystem mit Hilfe von Zusatzmuskeln geregelt werden. Die zweite Frequenz hängt vermutlich von den elastischen Eigenschaften der Thoraxkapsel ab und kann deshalb wahrscheinlich nur auftreten, wenn die indirekten Flugmuskeln momentan erschlaffen. Sie wird wahrscheinlich durch Muskeln verändert, die die Steifheit der Thoraxkapsel regulieren. Sie liegt meist im kHz-Bereich. Da ihre Phase in jedem halben Flügelschlag neu gesetzt wird, zeigt die Analyse im Sonagraph (Fourier Analyse) nur Harmonische der Flügelschlagfrequenz. Die Bedeutung der zweiten Frequenz für das Verhalten der Fliegen ist unbekannt. In der Kommunikation der Bienen könnte sie eine wichtige Rolle spielen, da die einzigen bisher bekannten Vibrationsrezeptoren, die Subgenualorgane, ihr Empfindlichkeitsmaximum bei 2,5 kHz haben.

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Supported by the Deutsche Forschungsgemeinschaft, and grant NB 03927 from the USPHS

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Herrn Professor H. Autrum zum 60. Geburtstag gewidmet.     

Directionality in the mechanical response to substrate vibration in a treehopper (Hemiptera: Membracidae: Umbonia crassicornis)

The use of substrate vibrations in communication and predator-prey interactions is widespread in arthropods. In many contexts, localization of the vibration source plays an important role. For small species on solid substrates, time and amplitude differences between receptors in different legs may be extremely small, and the mechanisms of vibration localization are unclear. Here we ask whether directional information is contained in the mechanical response of an insect's body to substrate vibration. Our study species was a membracid treehopper (Umbonia crassicornis) that communicates using bending waves in plant stems. We used a bending-wave simulator that allows precise control of the frequency, intensity and direction of the vibrational stimulus. With laser-Doppler vibrometry, we measured points on the substrate and on the insect's thorax and middle leg. Transfer functions showing the response of the body relative to the substrate revealed resonance at lower frequencies and attenuation at higher frequencies. There were two modes of vibration along the body's long axis, a translational and a rotational mode. Furthermore, the transfer functions measured on the body differed substantially depending on whether the stimulus originated in front of or behind the insect. Directional information is thus available in the mechanical response of the body of these insects to substrate vibration. These results suggest a vibration localization mechanism that could function at very small spatial scales.     

A method for two-dimensional characterization of animal vibrational signals transmitted along plant stems

Conventional approaches to measuring animal vibrational signals on plant stems use a single transducer to measure the amplitude of vibrations. Such an approach, however, will often underestimate the amplitude of bending waves traveling along the stem. This occurs because vibration transducers are maximally sensitive along a single axis, which may not correspond to the major axis of stem motion. Furthermore, stem motion may be more complex than that of a bending wave propagating along a single axis, and such motion cannot be described using a single transducer. Here, we describe a method for characterizing stem motion in two dimensions by processing the signals from two orthogonally positioned transducers. Viewed relative to a cross-sectional plane, a point on the stem surface moves in an ellipse at any one frequency, with the ellipse’s major axis corresponding to the maximum amplitude of vibration. The method outlined here measures the ellipse’s major and minor axes, and its angle of rotation relative to one of the transducers. We illustrate this method with measurements of stem motion during insect vibrational communication. It is likely the two-dimensional nature of stem motion is relevant to insect vibration perception, making this method a promising avenue for studies of plant-borne transmission. Electronic Supplementary Material The online version of this article () contains supplementary material, which is available to authorized users.     

How to obtain a bloodmeal without being eaten by a host: the case of poultry red mite, Dermanyssus gallinae

Abstract.  The behavioural responses of the poultry red mite, Dermanyssus gallinae , to three host-related stimuli, vibration, heat and CO₂, are studied in light and dark. Although D. gallinae usually feeds on host-blood at night, it can also be day active after a few days of starvation. However, the immediate response of the mites to CO₂ in daylight is to freeze and remain motionless. Even with simultaneous presentation of an activating stimulus (heat), the mites freeze in response to CO₂. In the case of subsequent vibrations, they start moving but only for the duration of the vibrations. After 2 min, the freezing response disappears and the level of activity is significantly higher than for mites stimulated with vibrations alone. The freezing response is interpreted as a defence against being eaten by the host that apparently is close enough to breathe on the mites. At low-light intensities, where the birds are unable to see the mites, there is no freezing response, but only a synergistic effect of heat and vibration on the level of activity. The frequency of vibration most efficient in eliciting movement during the freezing response is found at 2 kHz with a threshold value of 35 µm s⁻¹ peak-peak, suggesting that D. gallinae is more sensitive to substrate-borne vibrations than some insects (e.g. honeybees) known to use vibrations for intraspecific communication.     

Regulation of the number of spiders participating in collective prey transport in the social spider Anelosimus eximius (Araneae, Theridiidae)

In an experimental study, mechanisms by which cooperative prey transport is achieved in social spiders were clarified. Factors that could influence the number of individuals that participate in prey transport (prey mass, length and vibration) were investigated. Results show that two factors are fundamental: the vibrations and the prey length. Prey mass did not seem to influence spiders' participation. Thus, the single fact that individuals respond locally to environmental stimuli (intensity of vibration, available site on the prey) explains how spiders cooperate and efficiently capture a wide range of prey types without complex communication systems.     

Acoustic Component and Social Context of the Wing Display of the Walnut Fly Rhagoletis juglandis

Courtship signaling via wing vibration, accompanied by sound production, has been reported in several species of tephritids. In this large family of flies, sound communication as well as complex courtship displays appears to be restricted to species with lekking mating systems (i.e., Mediterranean fruit fly, Anastrepha and Dacus species). In contrast, in tephritid species with resource-defense mating systems, such as species in the genus Rhagoletis, little or no courtship behavior, acoustical or otherwise, has been described. Wing displays in Rhagoletis species have been considered to play a visual role. This study describes a distinctive wing display performed by males of the walnut fly, Rhagoletis juglandis. Laboratory experiments and field observations demonstrate that the male wing display plays a role in courtship. We used sound and vibration detectors to record the signals produced by this wing display. Using a combination of techniques, we were able to record both the very low-frequency vibration and its accompanying airborne infrasound (12–22 Hz) produced by the males.     

Predatory bug <Emphasis Type="Italic">Picromerus bidens</Emphasis> communicates at different frequency levels

The Asopinae (Heteroptera: Pentatomidae) are a subfamily of stinkbugs with predaceous feeding habits and poorly understood communication systems. In this study we recorded vibratory signals emitted by Picromerus bidens L. on a non-resonant substrate and investigated their frequency characteristics. Males and females produced signals by vibration of the abdomen and tremulation. The female and male songs produced by abdominal vibrations showed gender-specific time structure. There were no differences in the temporal patterns of male or female tremulatory signals. The signals produced by abdominal vibrations were emitted below 600 Hz whereas tremulatory signals had frequency ranges extending up to 4 kHz. Spectra of male vibratory signals produced by abdominal vibrations contained different peaks, each of which may be dominant within the same song sequence. Males alternated with each other during production of rivalry signals, using different dominant frequency levels. We show that the vibratory song repertoire of P. bidens is broader than those of other predatory stinkbugs that have been investigated. The emission of vibrational signals with different dominant frequencies but the same production mechanism has not yet been described in heteropteran insects, and may facilitate location of individual sources of vibration within a group.     

Response characteristics of vibration-sensitive neurons in the midbrain of the grassfrog,Rana temporaria

Summary European grassfrogs (Rana temporaria) were stimulated with pulsed sinusoidal, vertical vibrations (10–300 Hz) and the responses of 46 single midbrain neurons were recorded in awake, immobilized animals.Most units (40) had simple V-shaped excitatory vibrational tuning curves. The distribution of best frequencies (BF's) was bimodal with peaks at 10 and 100 Hz and the thresholds ranged from 0.02 to 1.28cm/s² at the BF.Twenty-three neurons showed phasic-tonic and 11 neurons phasic responses. The dynamic range of seismic intensity for most neurons was 20–30 dB.In contrast to the sharp phase-locking in peripheral vibration-sensitive fibers, no phase-locking to the sinusoidal wave-form was seen in the midbrain neurons. The midbrain cells did not respond at low stimulus intensities (below 0.01–0.02 cm/s²) where a clear synchronization response occurs in saccular fibers.Six midbrain neurons had more complex response characteristics expressed by inhibition of their spontaneous activity by vibration or by bi-and trimodal sensory sensitivities.In conclusion, the vibration sensitive cells in the midbrain of the grassfrog can encode the frequency, intensity, onset and cessation of vibration stimuli. Seismic stimuli probably play a role in communication and detection of predators and the vibration-sensitive midbrain neurons may be involved in the central processing of such behaviorally significant stimuli.Abbreviation BF best frequency