Rohini Balakrishnan

Species concepts, species boundaries and species identification : A view from the tropics

The species has been treated as a fundamental unit in biology (Hull, 1977) and, more recently, in biodiversity conservation (Sites and Crandall, 1997). Almost all studies in biology, whether at the level of molecules, cells, individuals or populations, are typically referenced to the level of the species. In the field of conservation biology, assessments of biodiversity are made at the level of the species: typical criteria include species richness, numbers of endemic species, and the number or presence of endangered species in given areas (Myers et al. 2000). The accurate identification of species is crucial both to research in all areas of biology and to biodiversity conservation. It is therefore surprising that, in the field of systematics, species are currently used mostly as terminal taxa in the reconstruction of phylogenetic trees, whereas the methods by which they are delimited and identified receive scant attention (Wiens and Penkrot, 2002).

Song pattern recognition in the grasshopper Chorthippus biguttulus: the mechanism of syllable onset and offset detection.

Abstract. The male song of the duetting grasshopper Chorthippus biguttulus consists of syllables alternating with noisy pauses. The syllable-pause structure is important for song recognition by the female. Using playback experiments we investigated the mechanism by which intensity modulations within the song pattern are used to detect syllable onsets and offsets. We varied the relative onset level (level of the syllable beginning relative to the noisy pause) and the relative offset level (level of the noisy pause relative to the syllable end) independently in different experiments. For all females, an increase in intensity defining the syllable onset was necessary to evoke responses. Syllable offset cues were not always necessary: some females responded to continuous noise stimuli wherein only syllable onsets were marked by short pulses of high intensity. Those females that did not require syllable offset cues did not, however, lack a functional pause detection mechanism, since their responses to model songs containing silent pauses were restricted to a given range of pause durations. We propose that syllable-pause detection involves two independent processes: (1) syllable onset detection by a phasic neuronal unit that can be re-activated only after a short pause, and (2) the rejection of unacceptably long pauses by a second unit.

Acoustic communication in a duetting grasshopper: Receiver response variability, male strategies and signal design

In the duetting grasshopper Chorthippus biguttulus, a females decision to reply to a conspecific male is based on the evaluation of a number of features of the males song, which consists of uninterrupted syllables separated by pauses. Female responses are tuned to a restricted range of pause durations. However, males produce songs with noisy rather than silent pauses, which should make the measurement of pause durations more difficult for the female. We examined the adaptive value of these noisy pauses by testing female responses to (1) pairs of natural phrases, which differed only with respect to clear or noisy syllable pauses, and (2) synthetic phrases, in which the syllable onset accentuations and noise levels in the pauses were systematically varied. There was considerable variation between females, both in their preference for clear or noisy pauses in natural phrases, and in the optimal combinations of syllable onset accentuations and noise levels in pauses that they preferred in synthetic phrases. The response profiles of individual females were consistent. The experiments with synthetic phrases showed that, on average, females preferred more extreme values of syllable onset accentuations than were present in male songs. Noisy pauses increased the range of syllable pause durations accepted by females. The results suggest that noisy pauses could buffer signallers against the negative consequences of both signal degradation during transmission and extreme receiver choosiness.

Vertical stratification in an acoustically communicating ensiferan assemblage of a tropical evergreen forest in southern India

This study provides the first quantitative description of vertical stratification in calling heights of ensiferan species constituting most of the dry-season nocturnal acoustic community of an evergreen forest in Kudremukh National Park in south-west India. Calling heights of an average of 26 individuals of each of the 20 ensiferan species were measured and subjected to an analysis of variance to test for differences in mean calling height between species and a cluster analysis to check for the presence of discontinuous calling height layers. There were significant differences in mean calling heights between species. Calling heights of different gryllid and tettigoniid species ranged from the ground to the canopy. More gryllid than tettigoniid species occupied the ground and herb layer. Our study revealed vertical stratification of calling heights, with discrete layers corresponding to the canopy, understorey and the ground layer. These clusters emerged from the raw data of calling heights of individuals without a priori distinction of layers. We found no significant correlation between the calling heights of species and call features, including mean dominant frequency of narrow band calls, mean syllable rate and mean duty cycle.

A DIVERSITY OF SONGS AMONG MORPHOLOGICALLY INDISTINGUISHABLE KATYDIDS OF THE GENUS MECOPODA ORTHOPTERA: TETTIGONIIDAE) FROM SOUTHERN INDIA

ABSTRACT Five distinct song types of the genus Mecopoda were found in Southern India. Four were morphologically indistinguishable. Some of them were both sympatric and had synchronous breeding seasons. The songs of these five song types ranged from simple short chirps to highly complex songs with multiple components. The temporal patterns of the songs of the five song types were very distinctive, whereas their spectral features were similar. Component elements of the different songs were distinct despite overall similarity. The song types possibly represent sibling species.

Vocalizations of wild Asian elephants (Elephas maximus): Structural classification and social context

Elephants use vocalizations for both long and short distance communication. Whereas the acoustic repertoire of the African elephant (Loxodonta africana) has been extensively studied in its savannah habitat, very little is known about the structure and social context of the vocalizations of the Asian elephant (Elephas maximus), which is mostly found in forests. In this study, the vocal repertoire of wild Asian elephants in southern India was examined. The calls could be classified into four mutually exclusive categories, namely, trumpets, chirps, roars, and rumbles, based on quantitative analyses of their spectral and temporal features. One of the call types, the rumble, exhibited high structural diversity, particularly in the direction and extent of frequency modulation of calls. Juveniles produced three of the four call types, including trumpets, roars, and rumbles, in the context of play and distress. Adults produced trumpets and roars in the context of disturbance, aggression, and play. Chirps were typically produced in situations of confusion and alarm. Rumbles were used for contact calling within and among herds, by matriarchs to assemble the herd, in close-range social interactions, and during disturbance and aggression. Spectral and temporal features of the four call types were similar between Asian and African elephants.

Synchrony during acoustic interactions in the bushcricket Mecopoda ‘Chirper’ (Tettigoniidae:Orthoptera) is generated by a combination of chirp-by-chirp resetting and change in intrinsic chirp rate

In several bushcricket species, individual males synchronise their chirps during acoustic interactions. Synchrony is imperfect with the chirps of one male leading or lagging the other by a few milliseconds. Imperfect synchrony is believed to have evolved in response to female preferences for leading chirps. We investigated the mechanism underlying synchrony in the bushcricket species Mecopoda ‘Chirper’ from Southern India using playback experiments and simulations of pairwise interactions. We also investigated whether intrinsic chirp period is a good predictor of leading probability during interactions between males. The mechanism underlying synchrony in this species differs from previously reported mechanisms in that it involves both a change in the oscillator’s intrinsic rate and resetting on a chirp-by-chirp basis. The form of the phase response curve differs from those of previously reported firefly and bushcricket species including the closely related Malaysian species Mecopoda elongata. Simulations exploring oscillator properties showed that the outcome of pairwise interactions was independent of initial phase and alternation was not possible. Solo intrinsic chirp period was a relatively good predictor of leading probability. However, changing the intrinsic period during interactions could enable males with longer periods to lead during acoustic interactions.

Changing resonator geometry to boost sound power decouples size and song frequency in a small insect

Despite their small size, some insects, such as crickets, can produce high amplitude mating songs by rubbing their wings together. By exploiting structural resonance for sound radiation, crickets broadcast species-specific songs at a sharply tuned frequency. Such songs enhance the range of signal transmission, contain information about the signaler’s quality, and allow mate choice. The production of pure tones requires elaborate structural mechanisms that control and sustain resonance at the species-specific frequency. Tree crickets differ sharply from this scheme. Although they use a resonant system to produce sound, tree crickets can produce high amplitude songs at different frequencies, varying by as much as an octave. Based on an investigation of the driving mechanism and the resonant system, using laser Doppler vibrometry and finite element modeling, we show that it is the distinctive geometry of the crickets’ forewings (the resonant system) that is responsible for their capacity to vary frequency. The long, enlarged wings enable the production of high amplitude songs; however, as a mechanical consequence of the high aspect ratio, the resonant structures have multiple resonant modes that are similar in frequency. The drive produced by the singing apparatus cannot, therefore, be locked to a single frequency, and different resonant modes can easily be engaged, allowing individual males to vary the carrier frequency of their songs. Such flexibility in sound production, decoupling body size and song frequency, has important implications for conventional views of mate choice, and offers inspiration for the design of miniature, multifrequency, resonant acoustic radiators.

Ecology of acoustic signalling and the problem of masking interference in insects

The efficiency of long-distance acoustic signalling of insects in their natural habitat is constrained in several ways. Acoustic signals are not only subjected to changes imposed by the physical structure of the habitat such as attenuation and degradation but also to masking interference from co-occurring signals of other acoustically communicating species. Masking interference is likely to be a ubiquitous problem in multi-species assemblages, but successful communication in natural environments under noisy conditions suggests powerful strategies to deal with the detection and recognition of relevant signals. In this review we present recent work on the role of the habitat as a driving force in shaping insect signal structures. In the context of acoustic masking interference, we discuss the ecological niche concept and examine the role of acoustic resource partitioning in the temporal, spatial and spectral domains as sender strategies to counter masking. We then examine the efficacy of different receiver strategies: physiological mechanisms such as frequency tuning, spatial release from masking and gain control as useful strategies to counteract acoustic masking. We also review recent work on the effects of anthropogenic noise on insect acoustic communication and the importance of insect sounds as indicators of biodiversity and ecosystem health.

Females of a tree cricket prefer larger males but not the lower frequency male calls that indicate large body size

In the tree cricket Oecanthus henryi, females are attracted by male calls and can choose between males. To make a case for female choice based on male calls, it is necessary to examine male call variation in the field and identify repeatable call features that are reliable indicators of male size or symmetry. Female preference for these reliable call features and the underlying assumption behind this choice, female preference for larger males, also need to be examined. We found that females did prefer larger males during mating, as revealed by the longer mating durations and longer spermatophore retention times. We then examined the correlation between acoustic and morphological features and the repeatability of male calls in the field across two temporal scales, within and across nights. We found that carrier frequency was a reliable indicator of male size, with larger males calling at lower frequencies at a given temperature. Simultaneous playback of male calls differing in frequency, spanning the entire range of natural variation at a given temperature, revealed a lack of female preference for low carrier frequencies. The contrasting results between the phonotaxis and mating experiments may be because females are incapable of discriminating small differences in frequency or because the change in call carrier frequency with temperature renders this cue unreliable in tree crickets