Analysis of waving and sound‐production display in the ghost crab,Ocypode stimpsoni

The ghost crab Ocypode stimpsoni displays waving and sound production. Sounds are produced by thumping the sand substratum with the major cheliped, and two types of sounds can be discriminated; one with a low frequency of about 12 Hz, called rapping, and another with a higher frequency (about double), called quivering. In our observations, a sequence of waving and sound emission would sometimes terminate abruptly, or appear as independent components but the component order never changed. The most frequently observed patterns were “waving with rapping and quivering”;, “waving with quivering”; and “quivering only”; quivering sounds being involved in more than 80% displays. Quivering sometimes occurred immediately after crabs emerged from the burrow, or when they returned to the entrance after discarding an excavated sand mass. The occurrence frequency of waving and sounds, the wave amplitude, and the frequency of the sound increased when other crabs approached.     

Ontogenetic development of auditory sensitivity and sound production in the squeaker catfish <Emphasis Type="Italic">Synodontis schoutedeni</Emphasis>

Background  

Surveys of ontogenetic development of hearing and sound production in fish are scarce, and the ontogenetic development of acoustic communication has been investigated in only two fish species so far. Studies on the labyrinth fish Trichopsis vittata and the toadfish Halobatrachus didactylus show that the ability to detect conspecific sounds develops during growth. In otophysine fish, which are characterized by Weberian ossicles and improved hearing sensitivities, the ontogenetic development of sound communication has never been investigated. We analysed the ontogeny of the auditory sensitivity and vocalizations in the mochokid catfish Synodontis schoutedeni. Mochokid catfishes of the genus Synodontis are commonly called squeakers because they produce broadband stridulation sounds during abduction and adduction of pectoral fin spines. Fish from six different size groups - from 22 mm standard length to 126 mm - were studied. Hearing thresholds were measured between 50 Hz and 6 kHz using the auditory evoked potentials recording technique; stridulation sounds were recorded and their sound pressure levels determined. Finally, absolute sound power spectra were compared to auditory sensitivity curves within each size group.     

Sounds and sound production in fishes

The main information on the sounds and sound production in fishes is reviewed. The present systems of sound classification and specialized sound production in fishes with different taxonomic positions and ecology are described. The anatomy of sound generating organs is analyzed, and the mechanisms of production of different types of sounds (stridulation, drumming, cavitation, and percussion, as well as hydrodynamic, pneumatic, stringed, and respiratory sounds) are discussed. A brief characterization of the acoustic parameters of different sound types is given. Recent data on the anatomy and morphology of the sonic muscles (including their innervation, physiology, sexual dimorphism, and seasonal changes) are reviewed. The dynamics of the development of sound generating organs are described, and their capacity for sound production in the ontogeny of fishes is followed.     

Geographic trends in mangrove crab abundance in East Africa

The aim of this work was to determine the abundance of crabs inmangrove communities along a latitudinal gradient along the eastern coastof Africa from 4°S to 32°S. Surveys were made atMombasa (Kenya), Zanzibar (Tanzania), Maputo (Mozambique) and in theTranskei (South Africa). Crabs were estimated at three designated levelsin the mangroves by visual census using a common protocol, and numberswere converted to biomass.Even after standardising the selection of sites and methods of censusthere was still extensive variability in the data, emphasising the complexheterogeneity of mangrove ecosystems. Lunar phase (full versus new moonsprings) did not have a consistent effect on results, but shore height hadseveral effects. Total crab biomass was similar in the two lower shore strataexamined, but about twice as high at the top-Avicennia level. Theratio of grapsid biomass:ocypodid biomass also changed with height: fromnear unity in the lower mangrove, to 0.14 in the middle strata, but to 15at the top.There was no consistent latitudinal trend in total crab numbers, but totalcrab biomass increased from north to south. In addition there was aconsistent and marked change in the grapsid biomass:ocypodid biomassratio: this swung from 0.65 at Mombasa to 6.8 in the Transkei. This hasimplications for the transfer of primary production through the food chain. Grapsids are important macrophagous feeders on the leaves and other partsof mangroves, whereas ocypodids are microphagous deposit feeders.     

Anatomy of the stridulation apparatus of the beech leaf‐mining weevil and characterization of,and behavioral responses to,stridulation sounds

We investigated auditory signals and morphology of the stridulatory apparatus of the European beech leaf‐mining weevil, Orchestes fagi L. (Coleoptera: Curculionidae), an invasive herbivore now established in Nova Scotia, Canada, to determine their potential for enhancing survey tools to monitor the spread of the species in Canada. We recorded and described sounds produced by adult O. fagi, analyzed the morphology of the stridulatory mechanism for intersexual differences and asymmetry, and examined behavioral responses elicited in conspecifics by playback of stridulation recordings. Adult O. fagi produced sounds under three conditions: male in distress, female in distress, and male in the presence of female. Female distress chirps lasted significantly longer than male distress chirps and male chirps in the presence of females, but peak frequencies and mean number of chirps per s did not differ significantly among the three groups. Morphology of the stridulation structures in male and female O. fagi was compared using scanning electron microscopy. Orchestes fagi have an elytro‐tergal file‐ and scraper‐type sound production apparatus, through which sound is produced upon anterior motion of the abdomen. Female O. fagi have a ‘pars stridens’ that is longer and has more ridges than males. Width and number of ridges per length of pars stridens did not differ between the sexes. Evidence of asymmetry was found in male pars stridens, with the right side being longer than the left. Playback of recorded sounds to adult weevils suggests female O. fagi were repelled by sounds produced by distressed males.     

Acoustic studies on the mechanism of sound production in the mating songs of the melon fly,Dacus cucurbitae Coquillett (Diptera: Tephritidae)

The sound production mechanism in the male mating songs for the melon flyDacus cucurbitae Coquillett was acoustically investigated to determine whether sounds resulted from free wing-fanning or stridulation produced by contact between wing and abdominal pecten. Waveforms in the songs of normal (pecten-present) males showed more complex vibrations than those of pecten-removed males. The total harmonic distortion in normal songs was greater than that of the pecten-removed songs. Female (pecten-absent) songs showed different sound signatures on the oscillogram and frequency spectrum from normal male songs. The female songs had more harmonic components than the normal male songs. These results suggest that wing/pecten stridulation of normal male songs produces complicated wing oscillations with a small number of harmonics of near-periodic waveforms and a high harmonic distortion. They also suggest that the free wing-fanning observed during female and pecten-removed male songs produces simple and uniform wing oscillations with many harmonic contents of complex-periodic waveforms and low harmonic distortion. Despite the occurrence of some acoustic differences, pecten-removal did not positively influence the rate of copulation.     

Gastric teeth of some thoracotreme crabs and their contribution to the brachyuran phylogeny

The gastric teeth of three ocypodoid species were investigated using scanning electron microscopy, and the morphological results were discussed with respect to the known food preferences. The species were chosen in particular because of contrasting ideas about their relationships within the Thoracotremata. For the genera Heloecius, Dotilla, Mictyris, and “Uca” (s. str.), we find a specific correlation of the gastric teeth with the suspension feeding. The lateral gastric teeth of Uca have no prominent lateral teeth cusps, and most of their teeth surface consists of transverse comb‐like lamellae. However, this possible food adaptation does not exclude the usability of specific teeth characters to distinguish species of suspension feeders. The closer relationship of the Dotillidae to grapsoid lines of gecarcinid or sesarmid crabs suggested by molecular data is not supported by the gastric teeth. For the genus Ucides, we found several characters that distinguish Ucides from the remaining ocypodoid genera Heloecius, Dotilla, Mictyris, and “Uca.” In particular, the structures of the lateral and the dorsomedian teeth show some similarities to genera of the Gecarcinidae and Sesarmidae. Our results suggest that foregut characters can be used for phylogenetic analyses. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Intracolony vibroacoustic communication in social insects

Vibrations and sounds, collectively called vibroacoustics, play significant roles in intracolony communication in termites, social wasps, ants, and social bees. Modalities of vibroacoustic signal production include stridulation, gross body movements, wing movements, high-frequency muscle contractions without wing movements, and scraping mandibles or tapping body parts on resonant substrates. Vibroacoustic signals are perceived primarily via Johnston’s organs in the antennae and subgenual organs in the legs. Substrate vibrations predominate as vibroacoustic modalities, with only honey bees having been shown to be able to hear airborne sound. Vibroacoustic messages include alarm, recruitment, colony activation, larval provisioning cues, and food resource assessment. This review describes the modalities and their behavioral contexts rather than electrophysiological aspects, therefore placing emphasis on the adaptive roles of vibroacoustic communication. Although much vibroacoustics research has been done, numerous opportunities exist for continuations and new directions in vibroacoustics research.     

Color change in exercising crabs: evidence for a hormone

Summary Three species of crabs exercised to fatigue showed a blanching and/or reddening of the body and legs. InUca pugilator this effect was due to white and red pigment dispersion in the leucophores and erythrophores, respectively, and a black pigment concentration in the melanophores. The pigment movements were induced by factor(s) present in the blood of exercisingUca; blood (hemolymph) removed from an exercised crab and injected into the isolated leg segment of another individual cause pigment movements similar to those seen in intact fatigued crabs. The blood of exercisedUca also caused similar chromatophore changes in isolated leg segments of the crabSesarma cinereum. The evidence suggests that blood-borne factor(s) related or identical to chromatophorotropins are released during vigorous exercise in crabs. We speculate that the effects of these exercise factor(s) are secondary to possible effects on carbohydrate and lipid metabolism associated with exercise.     

Acoustic defence in an insect: characteristics of defensive stridulation and differences between the sexes in the tettigoniid Poecilimon ornatus (Schmidt 1850)

Many insects exhibit secondary defence mechanisms upon contact with a predator, such as defensive sound production or regurgitation of gut contents. In the tettigoniid Poecilimon ornatus, both males and females are capable of sound production and of regurgitation. However, wing stridulatory structures for intraspecific acoustic communication evolved independently in males and females, and may result in different defence sounds. Here we investigate in P. ornatus whether secondary defence behaviours, in particular defence sounds, show sex-specific differences. The male defence sound differs significantly from the male calling song in that it has a longer syllable duration and a higher number of impulses per syllable. In females, the defence sound syllables are also significantly longer than the syllables of their response song to the male calling song. In addition, the acoustic disturbance stridulation differs notably between females and males as both sexes exhibit different temporal patterns of the defence sound. Furthermore, males use defence sounds more often than females. The higher proportion of male disturbance stridulation is consistent with a male-biased predation risk during calling and phonotactic behaviour. The temporal structures of the female and male defence sounds support a deimatic function of the startling sound in both females and males, rather than an adaptation for a particular temporal pattern. Independently of the clear differences in sound defence, no difference in regurgitation of gut content occurs between the sexes.     

The complex stridulatory behavior of the cricket Eneoptera guyanensis Chopard (Orthoptera: Grylloidea: Eneopterinae)

Crickets produce stridulated sounds by rubbing their forewings together. The calling song of the cricket species Eneoptera guyanensis Chopard, 1931 alternates two song sections, at low and high dominant frequencies, corresponding to two distinct sections of the stridulatory file. In the present study we address the complex acoustic behavior of E. guyanensis by integrating information on the peculiar morphology of the stridulatory file, the acoustic analysis of its calling song and the forewing movements during sound production. The results show that even if E. guyanensis matches the normal cricket functioning for syllable production, the stridulation involves two different closing movements, corresponding to two types of syllables, allowing the plectrum to hit alternately each differentiated section of the file. Transition syllables combine high and low frequencies and are emitted by a complete forewing closure over the whole file. The double-teeth section of the stridulatory file may be used as a multiplier for the song frequency because of the morphological multiplication due to the double teeth, but also because of an increase of wing velocity when this file section is used. According to available phylogenetic and acoustic data, this complex stridulation may have evolved in a two-step process.     

Auditory information processing in stridulating grasshoppers: tympanic membrane vibrations and neurophysiology

During stridulation in the gomphocerine grasshopper Omocestus viridulus the leg movements, sound pattern and either summed auditory nerve activity or single interneuron activity were recorded. Simultaneous laser interferometric and vibrometric measurements of the displacement and velocity of the tympanic membrane were performed at the pyriform vesicle (d-receptor group). Slow displacements of the tympanic membrane occur in phase with the ventilatory and stridulatory rhythm and reach 10 m_peak-peak and 1–3 m_peak-peak in amplitude, respectively. Additionally, the tympanic membrane oscillates maximally in the range 5–10 kHz. These high-frequency oscillations are due to sound production and motor activity and correspond in amplitude to oscillations evoked by sound pressures of 90-dB SPL. They activate the auditory receptors during most of the stridulatory cycle even during mute stridulation. Only at the lower reversal point of the leg movement are membrane vibrations and receptor activity at a minimum. As a consequence the response of receptors and interneurons to auditory stimuli are generally impaired and an auditory response of receptors and interneurons can be elicited only during a short period at the lower reversal point. Although in this phase of the stridulatory cycle auditory sensitivity is present, males do not show phonotactic responses towards female songs during ongoing own stridulation.     

UNUSUAL SONG AND RESPONSES IN A CHAFFINCH FRINGILLA COELEBS

ABSTRACT

Acoustic calling and the mechanisms of its production are described for the ghost crabs Ocypode jousseaumei, O. platytarsus, O. rotundata and O. saratan from Oman. Sounds generated by the major cheliped suggest the existence of species specific differences, predominantly in the temporal properties of rapping and in the physical structure of stridulation. The spectral energy of all sounds is concentrated at frequencies below 4 kHz. O. rotundata and O. saratan build sand pyramids adjacent to their courtship burrows and are considerably less acoustically active than O. jousseaumei and O. platytarsus that do not build pyramids, suggesting that the visual signal is substituted by an auditory one.     

Post‐Contest Stridulation Used Exclusively as a Victory Display in Mangrove Crabs

Victory or triumph display is a post‐contest signal, performed only by winners and not by losers. While much is unknown about its function, there is mounting evidence that victory displays are widespread among animals. However, evidence remains anecdotal in crabs. Sesarmid crabs belonging to the genera Parasesarma and Perisesarma are known to have characteristic stridulatory structures on their chelipeds. In Perisesarma eumolpe, a mangrove crab, stridulation has been anecdotally purported as a triumph display. We examined whether stridulation in P. eumolpe is a victory display and the factors affecting it by staging 17 contest trials among males and investigating the factors influencing stridulations and fight outcome in 55 fights. Using generalised linear mixed‐effects models, we find that stridulations were generally performed by winners and after fights, especially when the fights were intense. In addition, stridulation was only observed in the context of a contest, never before or outside of it. Stridulation in P. eumolpe is likely a victory display, and, unlike other forms of victory display described for other species, it appears exclusively used for asserting victory.     

Visual and Acoustical Signalling during Courtship by Fiddler Crabs (Genus Uca)

Male fiddler crabs (Genus Uca) employ both visual and acousticalsignals to attract females for mating. In U. pugilator and severalother American species, the males attract females during theday first by waving, then by producing sounds just within theirburrows. At night, the males produce sounds at low rates, butwhen touched by a female, they increase their rate of soundproduction. In the European species, U. tangeri, many elements of courtshipare similar to those in U. pugilator, but two types of soundsare produced. One of these, the short drumwhirl, appears tosubstitute for waving when the male is temporarily obscuredfrom the female during his diurnal courtship activities. Thelong drumwhirl is used under different circumstances. The acoustical responses of a male to a female influence thecourtship behavior of other males in the area. When sounds fromstimulated males are played back to test males during the day,their lates of waving increase. At night, the playbacks elicitincreases in rates of sound production. The influence of tidal oscillations, temperature, and lightcycles on the behavior of males is discussed. Courtship activities of aquatic crabs are compared to thoseof terrestrial Brachyura. In aquatic forms, courtship may beabsent or, if present, does not involve elaborate signallingby the male. Chemical or visual cues at close range are themost important stimuli. In several genera of terrestrial crabs,visual signalling for prolonged periods is common, and soundsare often emitted by males to "call" females from their burrowsto the surface for mating. Some of the factors that may accountfor differences in courtship activities in aquatic and terrestrialspecies are discussed.     

Locomotor Activity of the Fiddler Crab,Uca subcylindrica (Stimpson), under Artificial Illumination

Specimens of the fiddler crab Uca subcylindrica (Stimpson) were captured in south Texas (USA) for locomotor rhythm studies. Actographic data were analyzed using Tau? sofware. Under constant illumination (LL) and darkness (DD), the semiterrestrial crabs express a circadian rhythm of locomotion. When exposed to illumination/darkness cycles (LD_12:12 or LD_14:10), their bouts of activity are entrained to the photoperiod. In LD, activity is generally bimodal with an initial burst about 0.5 h after illumination. A second burst of activity begins 1 to 2 h before the end of illumination. When transferred from LD to LL, a locomotor rhythm with an average period of 24.6 ± 1.0 h (n = 19) is expressed in 89 percent of the crabs. On the other hand, when placed in DD after LD (n = 8), the crabs are either arrythmic or weakly rhythmic (period = 23.8 ± 0.2 h; n = 2). If the onset of illumination is advanced by 6 h, a period of less than 24.0 h is detected in the actogram. If the onset of illumination is delayed by 6 h, a locomotor rhythm with a period greater than 24.0 h appears. The locomotor behavior of this species of fiddler crab, Uca subcylindrica, is not related to the tidal rhythmicities seen in other members of the genus Uca. Rather, it has strong circadian components.     

Pharmacological brain stimulation releases elaborate stridulatory behaviour in gomphocerine grasshoppers – conclusions for the organization of the central nervous control

Grasshoppers produce a variety of sounds generated by complex movements of the hindlegs. Stridulation, performed in the context of partner finding, mating and rivalry, can be released by pressure injection of cholinergic agonists into the protocerebrum. Particularly stimulation with muscarinic agonists induced long-lasting stridulation that resembled the natural behaviour to an astonishing degree, not only with respect to their temporal structure and right/left coordination, but also to changes in the song sequences according to the progress of courtship stridulation, even including accessory movements of other parts of the body. According to the complexity of their stridulatory behaviour ten gomphocerine species were chosen for this comparative study. The results indicate that the protocerebrum fulfils two important tasks in the control of stridulation: (1) it integrates sensory input relevant to stridulation that represents a certain behavioural situation and internal state of arousal, and (2) it selectively activates and deactivates the thoracic networks that generate the appropriate movement and sound patterns. With the knowledge of the natural behaviour and the accessibility to pharmacological and electrophysiological studies, the cephalic control system for stridulation in grasshoppers appears to be a suitable model for how the brain selects and controls appropriate behaviours for a given situation.     

Calling Behavior and Morphology of the Stridulatory Apparatus in Neotropical Grasshoppers (Aleuas Stål; Orthoptera)

The sound production and sound producing apparatus of two species of grasshoppers, A. lineatus Stål, 1878 and A. vitticollis Stål, 1878 (Orthoptera, Acrididae, Copiocerinae) have been studied. For both species oscillograms and other physical data on sounds are presented for the first time. The sound producing behavior is described and the stridulation apparatus mechanism is illustrated. In the case of A. vitticollis these data are the first known on its sound production and general acoustic behavior.     

Function of Sound Production by the Skunk Loach Botia horae (Pisces,Cobitidae)

In agonistic interactions with conspecifics, skunk loaches (Botia horae) typically exhibit lateral displays, attacks, and produce a sound (a “click”). Four experiments were conducted to determine the functional significance of the click sound. Behaviors of resident fish were monitored while characteristics of their shelter and the size or type of intruder fish introduced were varied. Manipulations in the four experiments involved: 1) the configuration of the available shelter; 2) the size or recent history of intruder fish; 3) playbacks of “click” sounds or aquarium noise; or 4) muting of resident fish. Lateral displays, attacks, and sounds produced by residents were monitored, and the responses of residents were analyzed in terms of the functional significance of “click” sounds to juvenile skunk loaches.     

Contribution to the study of acoustic communication in two Belgian river bullheads (<Emphasis Type="Italic">Cottus rhenanus and C. perifretum</Emphasis>) with further insight into the sound-producing mechanism

Background

The freshwater sculpins (genus Cottus) are small, bottom-living fishes widely distributed in North America and Europe. The taxonomy of European species has remained unresolved for a long time due to the overlap of morphological characters. Sound production has already been documented in some cottid representatives, with sounds being involved in courtship and agonistic interactions. Although the movements associated with sound production have been observed, the underlying mechanism remains incomplete. Here, we focus on two closely related species from Belgium: C. rhenanus and C. perifretum. This study aims 1) to record and to compare acoustic communication in both species, 2) to give further insight into the sound-producing mechanism and 3) to look for new morphological traits allowing species differentiation.

Results

Both Cottus species produce multiple-pulsed agonistic sounds using a similar acoustic pattern: the first interpulse duration is always longer, making the first pulse unit distinct from the others. Recording sound production and hearing abilities showed a clear relationship between the sound spectra and auditory thresholds in both species: the peak frequencies of calls are around 150 Hz, which corresponds to their best hearing sensitivity. However, it appears that these fishes could not hear acoustic signals produced by conspecifics in their noisy habitat considering their hearing threshold expressed as sound pressure (~ 125 dB re 1 μPa). High-speed video recordings highlighted that each sound is produced during a complete back and forth movement of the pectoral girdle.

Conclusions

Both Cottus species use an acoustic pattern that remained conserved during species diversification. Surprisingly, calls do not seem to have a communicative function. On the other hand, fish could detect substrate vibrations resulting from movements carried out during sound production. Similarities in temporal and spectral characteristics also suggest that both species share a common sound-producing mechanism, likely based on pectoral girdle vibrations. From a morphological point of view, only the shape of the spinelike scales covering the body allows species differentiation.