The sounds of silence: cessation of singing and song pausing are ultrasound-induced acoustic startle behaviors in the katydid Neoconocephalus ensiger (Orthoptera; Tettigoniidae)

Previous studies of acoustic startle in insects have dealt with behavioral and/or neural mechanisms employed in evading aerially hawking, echolocating bats; however, insects also face terrestrial predators. Here we describe an acoustic startle response of the nocturnal katydid, Neoconocephalus ensiger. Stridulating males disturbed in the field perform obvious anti-predatory behaviors – cessation of singing, freezing, jumping, and evasive flight. Under controlled laboratory conditions we found that cessation of singing and song pausing are ultrasound-specific behaviors: when stimulated with pulsed ultrasound (20–100 kHz), but not audio-sound (<20 kHz), males cease mate calling or insert pauses in their song. A second factor influencing acoustic startle is the phase of stimulation: an acoustic startle response occurs only when the pulse of ultrasound arrives during the window of silence between stridulatory syllables. The average startle threshold and response latency was 70 ± 5 dB SPL and 34.2 ± 6.0 ms, respectively. N. ensiger is particularly useful for examining acoustic startle responses of non-flying insects because (1) its calling song is broadband and contains ultrasound, thus the possibility exists of confusion over the biological meaning of ultrasound, and (2) this species shows the classic bat-avoidance response while flying, so a direct comparison between two types of acoustic startle is possible within the same species. Accepted: 6 November 1999     

Structures on the antennal flagellum of a caddisfly,frenesia missa (Trichoptera,Limnephilidae)

Non-innervated macrotrichia and microtrichia, thick-walled chemoreceptors and three kinds of thin-walled chemoreceptors are present on the antennal flagellum of Frenesia missa. One of the thin-walled receptors, the plate organ, is of a type not previously recorded for any insect. About four times as many plate organs are present on the flagellum of the male as on that of the female. They occur also on the maxillary and labial palps.     

Molecular Phylogenetics of the Genus Neoconocephalus (Orthoptera,Tettigoniidae) and the Evolution of Temperate Life Histories

Background

The katydid genus Neoconocephalus (25+ species) has a prominent acoustic communication system and occurs in large parts of the Neotropics and Nearctic. This group has been subject of numerous behavioral, physiological, and evolutionary studies of its acoustic communication system. Two distinct life histories occur in this group: The tropical life history incorporates multiple generations/year and direct egg development without environmental triggers. Temperate life history is characterized by overwintering in the egg stage, cold trigger of egg development, and one generation/year. This study reconstructs the phylogenetic relationships within the genus to (1) determine the evolutionary history of the temperate life history, and (2) to support comparative studies of evolutionary and physiological problems in this genus.

Methodology/Principal Findings

We used Amplified Fragment Length Polymorphisms (AFLP), and sequences of two nuclear loci and one mitochondrial locus to reconstruct phylogenetic relationships. The analysis included 17 ingroup and two outgroup species. AFLP and mitochondrial data provided resolution at the species level while the two nuclear loci revealed only deeper nodes. The data sets were combined in a super-matrix to estimate a total evidence tree. Seven of the temperate species form a monophyletic group; however, three more temperate species were placed as siblings of tropical species.

Conclusions/Significance

Our analyses support the reliability of the current taxonomic treatment of the Neoconocephalus fauna of Caribbean, Central, and North America. Ancestral state reconstruction of life history traits was not conclusive, however at least four transitions between life histories occurred among our sample of species. The proposed phylogeny will strengthen conclusions from comparative work in this group.     

Reliable acoustic cues for female mate preference in a katydid (Scudderia curvicauda, Orthoptera: Tettigoniidae)

The call of male Scudderia curvicauda (Orthoptera: Tettigoniidae)consists of a series of phrases, and each phrase contains syllables.Females respond to the male signal with ticks that follow malephrases after a specific period of time. Pair formation takesplace after males locate the female using her response sounds.Repeated recordings of males revealed that the average numberof syllables produced per phrase was a table, within-male parameterand that this parameter was a reliable predictor of male size(pronotum length). Thus, phrase length could be a reliable cueby which females evaluate males. We presented virgin femaleswith a sequential choice of two tape-recorded male calls thatdiffered only in the mean number of syllables produced per phrase.Two different playback tapes were used, and each female wastested on each of 5 consecutive days with the same playbacktape. Females responded more often and with a greater numberof ticks to calls containing more syllables per phrase, andthis preference was maintained throughout the testing period.Male size was a poor predictor of the size of the spermatophorefood-gift produced by the male; therefore, females are probablynot selecting males for this attribute. For one of the playbacktapes, there was a significant increase in female responsivenessover several playback trials, suggesting that females may employa falling-threshold tactic with respect to mate preference.     

Evolution of Novel Signal Traits in the Absence of Female Preferences in Neoconocephalus Katydids (Orthoptera,Tettigoniidae)

Background Significance

Communication signals that function to bring together the sexes are important for maintaining reproductive isolation in many taxa. Changes in male calls are often attributed to sexual selection, in which female preferences initiate signal divergence. Natural selection can also influence signal traits if calls attract predators or parasitoids, or if calling is energetically costly. Neutral evolution is often neglected in the context of acoustic communication.

Methodology/Principal Findings

We describe a signal trait that appears to have evolved in the absence of either sexual or natural selection. In the katydid genus Neoconocephalus, calls with a derived pattern in which pulses are grouped into pairs have evolved five times independently. We have previously shown that in three of these species, females require the double pulse pattern for call recognition, and hence the recognition system of the females is also in a derived state. Here we describe the remaining two species and find that although males produce the derived call pattern, females use the ancestral recognition mechanism in which no pulse pattern is required. Females respond equally well to the single and double pulse calls, indicating that the derived trait is selectively neutral in the context of mate recognition.

Conclusions/Significance

These results suggest that 1) neutral changes in signal traits could be important in the diversification of communication systems, and 2) males rather than females may be responsible for initiating signal divergence.     

Ecology and sound signaling of relict katydid Deracantha onos (Pallas, 1772) (Orthoptera, Tettigoniidae, Bradyporinae)

Life history and acoustic signaling of a Central Asian species Deracantha onos (Pall.) are described. Males and females produce calling signals and protest sounds. For the first time, electronograms of stridulatory files, oscillograms, and frequency spectra of some acoustic signals are given.     

Ultrastructure and electrophysiology of thermosensitive sensilla coeloconica in a tropical katydid of the genus Mecopoda (Orthoptera,Tettigoniidae)

In many acoustic insects, mate finding and mate choice are primarily based on acoustic signals. In several species with high-intensity calling songs, such as the studied katydid Mecopoda sp., males exhibit an increase in their thoracic temperature during singing, which is linearly correlated with the amount of energy invested in song production. If this increased body temperature is used by females as an additional cue to assess the male's quality during mate choice, as has been recently hypothesized (“hot-male” hypothesis), thermosensory structures would be required to evaluate this cue. In the present study, therefore, we investigated the ultrastructure and physiology of thermosensitive sensilla coeloconica on the antennal flagella of Mecopoda sp. using a combination of electron microscopy and electrophysiological recording techniques.We could identify three distinct types of sensilla coeloconica based on differences in the number and branching pattern of their dendrites. Physiological recordings revealed the innervation by antagonistically responding thermoreceptors (cold and warm) and bimodal hygro-/thermoreceptors (moist or dry) in various combinations. Our findings indicate that Mecopoda sp. females are capable of detecting a singing male from distances of at least several centimetres solely by assessing thermal cues.     

Reverse stridulatory wing motion produces highly resonant calls in a neotropical katydid (Orthoptera: Tettigoniidae: Pseudophyllinae)

This paper describes the biomechanics of an unusual form of wing stridulation in katydids, termed here 'reverse stridulation'. Male crickets and katydids produced sound to attract females by rubbing their forewings together. One of the wings bears a vein ventrally modified with teeth (a file), while the other harbours a scraper on its anal edge. The wings open and close in rhythmic cycles, but sound is usually produced during the closing phase as the scraper moves along the file. Scraper-tooth strikes create vibrations that are subsequently amplified by wing cells specialised in sound radiation. The sound produced is either resonant (pure tone) or non-resonant (broadband); these two forms vary across species, but resonant requires complex wing mechanics. Using a sensitive optical diode and high-speed video to examine wing motion, and Laser Doppler Vibrometry (LDV) to study wing resonances, I describe the mechanics of stridulation used by males of the neotropical katydid Ischnomela gracilis (Pseudophyllinae). Males sing with a pure tone at ca.15 kHz and, in contrast to most Ensifera using wing stridulation, produce sound during the opening phase of the wings. The stridulatory file exhibits evident adaptations for such reverse scraper motion. LDV recordings show that the wing cells resonate sharply at ca. 15 kHz. Recordings of wing motion suggest that during the opening phase, the scraper strikes nearly 15,000 teeth/s. Therefore, the song of this species is produced by resonance. The implications of such adaptations (reverse motion, file morphology, and wing resonance) are discussed.     

Stridulatory movements in eight species of Neoconocephalus (Tettigoniidae)

High-speed sound-synchronized photography showed details of stridulatory movements and their relation to the sounds produced. Closing movements were more prolonged than opening movements and generally produced more intense sounds. Wing closures varied in rate from 10 to 250/sec. Four species had unpredicted complexities in their stridulatory movements: one silently snapped its wings shut after each acoustically effective stroke; two alternated long and short wing closures; and one made repeated groups of 4 to 9 progressively changing closures.     

Colour polymorphism in Ephippiger ephippiger (Orthoptera, Tettigoniidae)

Colour in the Ephippiger ephippiger complex varies with the geographical distribution and with rearing density. In the W European subspecies E. e. vitium , wild insects from northern and eastern parts of the range tend to have green tergites with relatively inconspicuous banding, whereas those from S France have conspicuously pale banded dark-green or brown tergites. These forms represent the ends of a continuum and there is no zone of distinction. However, the E Pyrenean subspecies, E. e. cunii , lacks the broad banding of its neighbouring E. e. vitium. The tergites of laboratory cultured insects of these stocks when reared in isolation are distinctly green but when reared together become progressively brown. The darkness of the brown is density dependant, so that at high densities the tergites may become almost black. These colour changes are irreversible. Adult insects also become darker with sexual maturity and after contact on mating. Wild-caught E. e. ephippiger from Italy and Yugoslavia have green tergites without obvious banding. Colour changes have not been found or induced in this subspecies. The significance of the colour change is considered and although firm data are wanting, circumstantial evidence suggests a deliberate shift to an aposematic or pseudoaposematic colour pattern.     

Sense organs on the antennal flagellum of psocids (Insecta,Psocoptera)

The sense organs on the antennal flagella of five species of winged psocids belonging to two families of Psocoptera, Psocidae and Leptopsocidae, have been examined. All agree in possessing tactile hairs, thick-walled chemoreceptors and long, porous chemoreceptors. Thin-walled chemoreceptors were identified in all species except Metylophorous novaescotiae. Coeloconic chemoreceptors were present in all species except Echmepteryx hageni. Campaniform sense organs were found only in Metylophorus novaescotiae and Psocus leidyi.     

Sense organs on the antennal flagellum of mayflies (Ephemeroptera)

A few very small sensory pegs with the characteristics of chemoreceptors are present in both males and females on the antennal flagellum of the mayflies, Stenacron interpunctatum (Say)

1 Stenacron interpunctatum (Say) is Stenonema interpunctatum (Say) of earlier literature.

and Cloeon sp. They are situated near the proximal end of the flagellum on its ventro-lateral surface and have escaped the attention of earlier investigators.     

Structure of the stridulation ofOrchelimum (Orthoptera: Tettigoniidae)

Summary The structure of the stridulation was investigated by re-playing tape-recordings at very slow speed. The findings were corrobrated by sonograms and mingograms.The central part of the song is the ripple, a fast succession of syllables around which isolated syllables (clicks) are distributed according to species and circumstances. The rate of syllables in the ripple is a linear function of temperature.A quantitative expression for the stridulatory activity is the actual number of syllables per time unit, including pauses. By changes in the combination of elements, at leastO. agile is able to increase the output of syllables four to six times.This work was made possible by a grant from the Carlsberg Foundation to whom my most sincere thanks are due. —As I do not have advanced equipment for sound analysis, I am very much indebted to Dr. Bondesen and cand. sci. Poul Hansen, Bioakustisk Laboratorium, Naturhistorisk Museum, Aarhus, Denmark, and Mr. W.B. Broughton and Dr. M. Samways, Animal Acoustic Unit, City of London Polytechnic, London, for analysing part of the material by sonograms and mingograms. The very valuable help of Dr. Th.J. Walker, University of Florida, Gainesville, Florida, in identifying the species is gratefully acknowledged. For friendly discussions and linguistic corrections my best thanks are due to H.T. Evans and F.D.S. Evans.     

Sense organs on the antennal flagellum of the human louse,Pediculus humanus (Anoplura)

Tactile hairs are present on all three subsegments of the antennal flagellum of the human louse. There is, in addition, a single chemoreceptor (tuft organ) on subsegment 2 and 12 or 13 chemoreceptors (one tuft organ, two pore organs and nine or ten pegs) on subsegment 3. The cuticle surrounding the bases of the pegs at the tip of the antenna is unusual in that parts of it are perforated by many fine pores. This cuticle is underlain by a thin layer of dendrites. This region may also have a chemoreceptor function.     

Ultrasonic startle behavior in bushcrickets (Orthoptera; Tettigoniidae)

1. In the present work, we show that in flight, bushcrickets not previously known to respond to ultrasound alter their flight course in response to ultrasonic stimuli. Such stimuli elicit in flying Neoconocephalus ensiger an extension of the front and middle legs along the body and a rapid closure of all 4 wings (Fig. 1). This is a short latency acoustic startle response to ultrasound, consistent with acoustic startle responses of other insects. 2. The percentage of trials on which acoustic startle responses were elicited was maximum (90%) for sound frequencies ranging from 25 to at least 60 kHz. No acoustic startle response was observed at frequencies of 5 or 10 kHz (Fig. 2). The threshold for the response was roughly 76 dB between 25 to 60 kHz (Fig. 2) and the behavioral latency was 45 ms (Fig. 3). Recordings from flight muscles show that they cease discharging during the acoustic startle response (Fig. 4). 3. The characteristics of the acoustic startle response match those of an auditory interneuron called the T-neuron. The frequency sensitivity of this neuron is greatest for sound frequencies ranging from 13 to 60 kHz (Fig. 6). Moreover, we found that the neuron produces many more spikes to ultrasound (30 kHz) of increasing intensities than to a conspecific communication sound, whose dominant frequency is 14 kHz (Fig. 7).     

Dispersal in a metapopulation of the bush cricket, Metrioptera bicolor (Orthoptera: Tettigoniidae)

1. I present a stochastic simulation model that describes individual movements of Metrioptera bicolor Philippi in a heterogeneous landscape, consisting of patches of suitable habitat surrounded by a matrix of unprofitable habitats. Although the model is parameterized with information about daily movement behaviour, it can generate spatially explicit predictions about inter-patch dispersal rates for much longer periods, e.g. one generation.
2. Long-term dispersal experiments were conducted to evaluate model predictions. Patch-specific emigration rates and the total distance moved by individuals could be predicted with satisfactory precision. Because of the stochastic nature of the model, it failed to predict which recipient patches emigrating individuals actually chose in a particular situation.
3. Spatially explicit simulations of the movement model were made for the whole natural distribution area of M. bicolor . The results suggest that emigration rates are negatively correlated with patch size. Local populations occurring on small patches may be more prone to extinction than those on large patches, by losing more emigrants than are compensated for by immigration.     

Sense organs on the antennal flagellum of two species of embioptera (Insecta)

Five different kinds of sense organs have been identified on the antennal flagellum of males of Ptilocerembia sp: (1) tactile hairs, (2) thick-walled chemoreceptors, (3) chemoreceptors with thick walls and many pores, (4) thin-walled chemoreceptors and (5) campaniform sense organs. The third type has not been reported previously for other species of insects and combines the characteristics of typical thick-walled and thin-walled chemoreceptors. The campaniform sense organs are situated in sets of three, 120° apart, on the periphery of the distal end of nearly every subsegment. They lie at the transition zone where the thicker cuticle of the subsegment changes to the thin membrane between subsegments.