Exploring the hidden landscape of female preferences for complex signals

A major challenge in evolutionary biology is explaining the origins of complex phenotypic diversity. In animal communication, complex signals may evolve from simpler signals because novel signal elements exploit preexisting biases in receivers’ sensory systems. Investigating the shape of female preference functions for novel signal characteristics is a powerful, but underutilized, method to describe the adaptive landscape potentially guiding complex signal evolution. We measured female preference functions for characteristics of acoustic appendages added to male calling songs in the grasshopper Chorthippus biguttulus, which naturally produces only simple songs. We discovered both hidden preferences for and biases against novel complex songs, and identified rules governing song attractiveness based on multiple characteristics of both the base song and appendage. The appendage's temporal position and duration were especially important: long appendages preceding the song often made songs less attractive, while following appendages were neutral or weakly attractive. Appendages had stronger effects on songs of shorter duration, but did not restore the attractiveness of very unattractive songs. We conclude that sensory biases favor, within predictable limits, the evolution of complex songs in grasshoppers. The function‐valued approach is an important tool in determining the generality of these limits in other taxa and signaling modalities.     

Noise affects the shape of female preference functions for acoustic signals

The shape of female mate preference functions influences the speed and direction of sexual signal evolution. However, the expression of female preferences is modulated by interactions between environmental conditions and the female's sensory processing system. Noise is an especially relevant environmental condition because it interferes directly with the neural processing of signals. Although noise is therefore likely a significant force in the evolution of communication systems, little is known about its effects on preference function shape. In the grasshopper Chorthippus biguttulus, female preferences for male calling song characteristics are likely to be affected by noise because its auditory system is sensitive to fine temporal details of songs. We measured female preference functions for variation in male song characteristics in several levels of masking noise and found strong effects of noise on preference function shape. The overall responsiveness to signals in noise generally decreased. Preference strength increased for some signal characteristics and decreased for others, largely corresponding to expectations based on neurophysiological studies of acoustic signal processing. These results suggest that different signal characteristics will be favored under different noise conditions, and thus that signal evolution may proceed differently depending on the extent and temporal patterning of environmental noise.     

A narrow hybrid zone between the grasshoppers Stenobothrus clavatus and Stenobothrus rubicundus (Orthoptera: Gomphocerinae): female preferences for courtship songs

Two grasshopper species, Stenobothrus rubicundus and Stenobothrus clavatus, were previously shown to hybridize in a narrow contact zone on Mount Tomaros in northern Greece. The species are characterized by complex and completely different courtship songs. In the present study, we investigated female preferences for the courtship songs of S. rubicundus, S. clavatus and hybrids in playback experiments. Playback of the courtship songs revealed assortative preferences in females of the parental species: they significantly more often preferred the songs of conspecific males. Hybrid females showed a lower selectivity than parental females, responding somewhat equally eager to playback of the songs of S. clavatus, S. rubicundus, and natural hybrid song, although less actively to the F₁ hybrid song. The results suggest that hybrid males may lose to males of parental species, whereas hybrid females would even have an advantage over parental females. Comparison of responses of females from allopatric populations and Mount Tomaros to different song types shows no evidence for reinforcement. Asymmetry found in female preferences may have implications for the structure of the hybrid zone. © 2013 The Linnean Society of London     

Acoustic signals in insects: A reproductive barrier and a taxonomic character

In singing insects, the song is an important component of the specific mate recognition system (SMRS). In communities of sympatric singing species, there is a partitioning of communication channels, the so-called “acoustic niches.” Within one community, the songs of different species always differ in temporal or frequency characters, i.e. occupy different acoustic niches. However, conspecific songs do not always act as an interspecific reproductive barrier, despite always being a SMRS component. The species that do not communicate acoustically due to allopatry, different timing of vocalization, inhabiting different biotopes, or unmatched food specializations can produce similar songs while forming reproductively isolated communities. Individuals of different sexes need not only to recognize a conspecific mate but also to evaluate its “quality.” The close-range signal (courtship song) provides more opportunities for choosing the “best” male than does the distant signal (calling song). In many species of Orthoptera, courtship includes not only acoustic but also vibrational, visual, chemical, and mechanical signals. An analysis of cricket songs showed the courtship songs to be on average more elaborate and variable than the calling songs. At the same time, due to the difference in mating behavior between the two groups, the acoustic component of courtship is used for mate quality evaluation to a greater extent in grasshoppers than in crickets. The courtship songs of grasshoppers are generally more elaborate in temporal structure than cricket songs; moreover, they may be accompanied by visual displays such as movements of various body parts. Thus, song evolution in grasshoppers is more strongly driven by sexual selection than that in crickets. According to the reinforcement hypothesis, the premating barrier between hybridizing species becomes stronger in response to reduced hybrid fitness. However, our behavioral experiments with two groups of hybridizing grasshopper species did not confirm the reinforcement hypothesis. We explain this, firstly, by a low level of genetic incompatibility between the hybridizing species and secondly, by high hybrid fitness when attracting a mate. A high competitive capability of hybrids may be accounted for by attractiveness of new elements in hybrid courtship songs. When we divide similar forms based on their songs, we in fact distinguish biological species using the criterion of their reproductive isolation. Acoustic differences between species are usually greater than morphological ones. Therefore, song analysis allows one to determine the real status of doubtful species-rank taxa, to distinguish species in a medley of sibling forms, and to reveal cryptic species in the cases when morphological studies fail to provide a univocal result. At the same time, songs are subject to intraspecific variation the range of which is different in different groups. Therefore, it is necessary to study which degree of difference corresponds to the species level before interpreting the status of some forms based on song comparisons. Besides, song similarities cannot indicate conspecificity of acoustically isolated forms; on the other hand, song differences between these forms prove that they are full-rank species.     

Broad Selectivity for Courtship Song in the Cricket Gryllus bimaculatus

Various characteristics of a long‐distance acoustic signal have been shown to vary to different degrees. It has been suggested that female preferences based on stable song parameters are stabilising or weakly directional, and preferences based on variable parameters are strongly directional. We tested this hypothesis based on a short‐distance signal (courtship song) produced by the field cricket, Gryllus bimaculatus. We studied the degree of variability of different courtship song parameters and the behavioural importance of several parameters using synthesised song models in playback experiments. We found that most of the courtship song elements of G. bimaculatus were quite variable (coefficient of variation, CV, in the range of 20–53%). The most variable parameter of the courtship song was the relative amplitude of two elements: high‐amplitude ticks and low‐amplitude pulses. Because songs containing only ticks (of rare occurrence) appeared to be more effective than songs with both ticks and pulses (of frequent occurrence), we consider female preferences to be directional. Alteration of less variable traits, such as the carrier frequency and duration of ticks (CV = 20–25%), had a different effect on female responsiveness. The synthesised songs with different carrier frequencies of ticks were as attractive to females as the positive control (courtship of muted males accompanied by playback of the recorded song). Altering the duration of ticks had a crucial effect on the female response rate, decreasing female responsiveness to the level observed in the negative control (courtship of muted males). Thus, we did not find a strong relationship between the variability of individual song parameters and their potential importance in song recognition and the evaluation of male quality. The partial inconsistency of our results with the data of other authors may be due to different patterns of past and current selection on long‐distance and short‐distance acoustic signals.     

Interspecific competition among grasshoppers and their effect on plant abundance in experimental field environments

Summary We tested whether grasshoppers in experimental field environments, i.e. cages (40×40 cm) placed on existing old field vegetation, (1) were limited in density by plant abundance and/or nitrogen content, (2) exhibited interspecific competition, and (3) altered the relative abundance of different plant species. We examined interactions among a pair of early season grasshopper species (May–June; Arphia conspersa and Pardalophora apiculata) and a late season pair (July–August; Melanoplus femur-rubrum and Melanoplus bivittatus). Each grasshopper species was placed in cages by itself and with another grasshopper species. Grasshoppers generally survived at higher density in fertilized cages and they reduced plant abundance relative to empty cages, suggesting that grasshoppers may be food limited at these densities. In unfertilized plots, early season grasshoppers preferred grasses (Schizachyrium scoparium and Poa pratensis) and favored the growth of forbs (especially Solidago spp.). However, late in summer, Melanoplus spp. preferred Solidago spp. and favored the growth of grasses.The pattern of grasshopper survivorship and plant reduction within these experimental environments provide preliminary support for some of the predictions of resource competition theory. Grasshoppers exhibited interspecific competition only if they significantly reduced plant biomass. If two species competed, a grasshopper species was eliminated only if the superior competitor, when living by itself, could reduce plant biomass to a significantly lower level than the inferior competitor. Competitors persisted only if they did not differ in their ability to reduce plant biomass or reduced the abundance of different plant species.     

Species Recognition During Substrate-Borne Communication in <Emphasis Type="Italic">Nezara viridula</Emphasis> (L.) (Pentatomidae: Heteroptera)

The information code in the temporal and spectral characteristics of the substrate-borne communication signals produced by insects has been primarily studied in insects in the suborder Auchenorrhyncha. In the present study we investigated which of the female calling song (FCS) parameters in Nezara viridula (L.) (Heteroptera, Pentatomidae) are essential for recognition by conspecific males. In playback experiments we measured male vibrational responsiveness to FCS signals varying in the durations of pulse trains and inter-pulse train intervals, repetition times, duty cycles, and dominant frequencies, and determined the preference range for each specific parameter. Males were able to distinguish songs of different temporal and frequency parameters and responded best to values characteristic of the song of conspecific females. Signal recognition is achieved on the basis of two temporal filters tuned to the durations of the pulse train and inter-pulse train interval. Males responded best to the dominant frequency characteristic of conspecific songs, which are tuned to the resonant properties of the herbaceous plants used for intraspecific signal transmission during communication.     

Grasshopper feeding rates, preferences, and growth on safflower

Laboratory experiments were conducted to measure the feeding rates, preferences, survival rates, maturation times, and weight gain of four common species of grasshoppers (Orthoptera: Acrididae) on four lines of safflower (Carthamus tinctorius). When safflower lines were presented individually, all were readily consumed although consumption differed significantly according to grasshopper species, age and sex, and to safflower line. Consumption was greatest on a safflower line devoid of spines. However, grasshoppers also fed readily on spiny lines, and results of experiments in which all safflower lines were presented together indicated preferences that did not necessarily correspond to the results of the single-line feeding trials. Camnula pellucida, a graminivorous species, did feed on safflower, but restricted feeding almost exclusively to stem cambium, whereas Melanoplus bivittatus, M. packardii and M. sanguinipes fed mainly on the heads, flowers and leaves. Highly significant differences in grasshopper maturation time and weight gain were apparent among grasshopper species, and among safflower lines. Seedtec-5, the line which was least digestible, least consumed and least preferred nevertheless yielded rapid development and the greatest body weights of grasshoppers. All species of Melanoplus developed more quickly and gained more weight on safflower than on wheat. One species, M. packardii, was more able to utilise safflower than its congeners, and may gain competitive advantage if safflower becomes widely grown.     

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

In the duetting grasshopper Chorthippus biguttulus, a female's decision to reply to a conspecific male is based on the evaluation of a number of features of the male's 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.     

Female House Wrens may Use a Low‐Amplitude Call as an Aggressive Signal

The majority of work on aggressive signaling has focused on male–male contests. Although females in many species compete over important resources, female signals are understudied. In house wrens (Troglodytes aedon), females compete with other females to protect nesting cavities and eggs. We suggest that a high‐pitched, low‐amplitude call, hereafter abbreviated as ‘HI’, may be an aggressive signal used by female house wrens. Using playback of simulated female intruders, we found that females used HI calls significantly more often during playback of female house wrens compared to playback of heterospecifics. Additionally, HI calls were given significantly more often in the minute preceding physical attack. In comparison, song rates did not predict future attacks. Finally, we present pilot data that suggest listening females may respond more aggressively to female playback containing HI calls compared to playback containing only songs. This suggests the reliability of HI calls could be maintained by a receiver retaliation rule. HI calls bear a striking resemblance to the low‐amplitude songs of many male songbirds, in terms of both acoustic structure and social context. This is one of the few reports of a putative low‐amplitude aggressive signal in a female songbird. However, the nature of female competition may make low‐amplitude signaling an underappreciated signaling form in female animals.     

ACOUSTIC COMMUNICATION DISTANCES OF A GOMPHOCERINE GRASSHOPPER

ABSTRACT

I studied in the laboratory and in the field the communication distance of the gomphocerine grasshopper Chorthippus biguttulus, which lives close to the ground in stony, short grass habitats. Using sound measurements and behavioural tests, I examined the frequency dependent attenuation and temporal degradation of the song pattern. Sound measurements (2–40 kHz) in the field revealed that excess attenuation increases with increasing frequency. Close to the ground, excess attenuation reaches 18 dB/m. On the basis of (i) excess attenuations, (ii) tympanic receptor threshold curves and (iii) song spectra, perception distances of female tympanic receptors for male songs and vice versa were calculated. Behavioural tests revealed that, especially in short and stony grass vegetation, the typical habitat of Ch. biguttulus, the temporal song pattern is strongly degraded. Thus, in the laboratory and in tall grass vegetation, sound attenuation was found to limit the communication distance. In contrast, in short grass vegetation, degradation of the temporal pattern was found to limit the communication distance of males listening to female songs. I argue that the exploitation of fast amplitude modulations for song recognition restricts the acoustic communication of grasshoppers to short distances (1–2.2 m). In this respect it seems adaptive to stridulate at low intensities.     

Song preference of female Bengalese finches as measured by operant conditioning

Male songbirds approach females by using their songs. Knowing what females perceive and prefer in male songs is an important aspect of understanding courtship behavior. Male Bengalese finches (Lonchura striata var. domestica) sing complex sequence patterns with their own variety of elements. We tested female preference with phonologically different songs and sequentially different songs using an operant selection task. In the first experiment we presented the father’s song and an unfamiliar song that were phonologically different. Females clearly preferred the father’s song over a novel song. In the second experiment, we used the father’s song and its element order reversed song to test female preference for the difference in element sequence. In the third experiment, we presented different sequence complexity songs edited from the same unfamiliar song elements to test female preference for complex sequence. Females did not show a significant preference in either the second or third experiment. Results show that female Bengalese finches discriminated the difference in song elements, preferred familiar songs, and did not show preference for difference of sequence. This study did not support evidence that female Bengalese finches prefer a complex sequence. However, in future research, we should carefully investigate female preference for choosing mates in a natural context.     

Effects of large herbivore grazing on grasshopper behaviour and abundance in a meadow steppe

1. Adaptive phenotypic plasticity has been a major subject in evolutionary ecology, but how a species' behaviour may respond to certain environmental change is still not clear. In grasslands worldwide, large herbivores are increasingly used as a tool for grazing management, and occur to interact with grasshoppers that dominate grassland insect communities. Previous studies have been well-documented about grazing effects on diversity and abundance of grasshoppers. Yet, how grazing may alter grasshopper behaviour, and potential effects on their abundance remains elusive. 2. We conducted a field experiment by manipulating grazing using sheep, cattle, and their mix to examine the behavioural responses and abundance of the grasshoppers (Euchorthippus unicolor) to grazing in a Leymus chinensis-dominated grassland. 3. Results showed that the grasshoppers spent less time on feeding and resting on grasses, but more time on switching and resting on forbs under cattle grazing and mixed grazing with cattle and sheep. In contrast, the grasshoppers spent more time on feeding but less time on switching and resting on forbs under sheep grazing. The behavioural changes were also potentially linked to grasshopper abundance in the context of grazing management. 4. The responses of grasshopper behaviour and abundance to grazing may be largely triggered by altered vegetation and microclimates. Such behavioural flexibility of grasshoppers must be considered when large herbivores are recognised as a management tool for influencing grasshopper abundance, and grazer species should be paid more attention both individually and jointly for better grassland conservation.     

Herbivore phenology can predict response to changes in plant quality by livestock grazing

Livestock grazing can have a strong impact on herbivore abundance, distribution and community. However, not all species of herbivores respond the same way to livestock grazing, and we still have a poor understanding of the underlying mechanisms driving these differential responses. Here, we investigate the effect of light intensity cattle grazing on the abundance of two grasshoppers (Euchorthippus cheui and E. unicolor) that co-occur in the same grasslands and feed on the same food plant (the dominant grass Leymus chinensis). The two grasshopper species differ in phenology so that their peak abundances are separated into early- and late-growing seasons. We used an exclosure experiment to monitor grasshopper abundance and food quality in the field under grazed and ungrazed conditions, and performed feeding trials to examine grasshopper preference for grazed or ungrazed food plants in the laboratory. We found that the nitrogen content of L. chinensis leaves continuously declined in the ungrazed areas, but was significantly enhanced by cattle grazing over the growing season. Cattle grazing facilitated the early-season grasshopper E. cheui, whereas it suppressed the late-season grasshopper E. unicolor. Moreover, feeding trials showed that E. cheui preferred L. chinensis from grazed plots, while E. unicolor preferred the leaves from ungrazed plots. We conclude that livestock grazing has opposite effects on the two grasshopper species, and that these effects may be driven by grazing-induced changes in plant nutrient content and the unique nutritional niches of the grasshoppers. These results suggest that insects that belong to the same guild can have opposite nutrient requirements, related to their distinct phenologies, and that this can ultimately affect their response to cattle grazing. Our results show that phenology may link insect physiological needs to local resource availabilities, and should be given more attention in future work on interactions between large herbivores and insects.     

Body size influences differently the detectabilities of colour morphs of cryptic prey

Body size and coloration may contribute to variation in performance and fitness among individuals; for example, by influencing vulnerability to predators. Yet, the combined effect of size and colour pattern on susceptibility to visual predators has received little attention, particularly in camouflaged prey. In the colour polymorphic pygmy grasshopper Tetrix subulata (Linnaeus, 1758), females are larger than males, although there is a size overlap between sexes. In the present study, we investigated how body size and colour morph influenced detection of these grasshoppers, and whether differences in protective value among morphs change with size. We conducted a computer‐based experiment and compared how human ‘predators’ detected images of large, intermediate or small grasshoppers belonging to black, grey or striped colour morphs when embedded in photographs of natural grasshopper habitats. We found that time to detection increased with decreasing size, that differences in time to detection of the black, grey and striped morphs depended differently on body size, and that no single morph provided superior or inferior protection in all three size classes. By comparing morph frequencies in samples of male and female grasshoppers from natural populations, we also examined whether the joint effects of size and colour morph on detection could explain evolutionary dynamics in the wild. Morph frequency differences between sexes were largely in accordance with expectations from the results of the detection experiment. The results of the present study demonstrate that body size and colour morph can interactively influence detection of camouflaged prey. This may contribute to the morph frequency differences between male and female pygmy grasshoppers in the wild. Such interactive effects may also influence the dynamics of colour polymorphisms, and contribute to the evolution of ontogenetic colour change and sexual dichromatism. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 112–122.     

‘Absolute steepness’ of ramps as an essential cue for auditory pattern recognition by a grasshopper (Orthoptera; Acrididae; Chorthippus biguttulus L.)

Male grasshoppers of the species Chorthippus biguttulus react to female songs with a characteristic turn towards the female. The probability of turning towards female song models was used to evaluate those parameters which are essential for a signal to be interpreted as female song.

1. The shape of sound pulses turned out to be most decisive; pulses with ramps rising gradually over 3 and more ms were efficient (Figs. 2, 3), whereas rectangularly modulated pulses evoked only weak responses and only when pulse intervals were between 3 and 5 ms (Fig. 2). The decline of a pulse did not influence its efficiency (Fig. 3). In particular, pulses with sudden onsets and gradual declines were as weakly effective as rectangularly modulated ones and thus remarkably less effective than pulses with ramp-like onsets (Fig. 4).
2. Intensity tuning curves suggest, that the absolute steepness of ramps (expressed as μbar/ms) is detected by the grasshopper nervous system (Figs. 6, 7), possibly by processing the delay in excitation onset of at least two receptor types differing in threshold sensitivity.
3. The sawtooth shape of pulses in female signals is suggested to be adaptive with respect to directional hearing.

     

Separate localization of sound recognizing and sound producing neural mechanisms in a grasshopper

Summary In the two acridid speciesChorthippus parallelus andCh. montanus, the sound template by which females recognize male song varies with temperature, as does the song itself. At relatively high temperatures the females respond best to simulated songs with high syllable frequencies, and at lower temperatures songs with lower syllable frequencies are preferred.The temperature around the supraesophageal and metathoracic ganglia of female grasshoppers was monitored by implanted thermocouples, and either the head or the thorax was warmed selectively while the animal was free to move (within the imits of the wires). Then simulations of the conspecific song varying in syllable frequency corresponding to different song temperatures were presented, and the stridulatory responses of the animals were observed.The results were as follows. 1. Song recognition (in particular, the position of the peak of the response curve) depended on the temperature of the head. 2. The rate of stridulatory hindleg movement was determined by the temperature of the thoracic ganglia.This result provides strong evidence against the genetic coupling hypothesis.     

Female recognition of trills in the male calling song of the field cricket,<Emphasis Type="Italic"> Teleogryllus taiwanemma</Emphasis>

The calling song of the field cricket, Teleogryllus taiwanemma, is usually considered to consist of sequences of separate chirps. However, sometimes it comprises a phrase of several chirps in a row, with one long chirp (chirp) and a few short chirps (trills). In this study, I compared the phrase containing only chirps with that containing both chirps and trills by analyzing male songs and conducting playback experiments of male songs to females. The song analyses showed significant differences between chirps and trills for all song parameters except bandwidth. To test whether female preference differed with respect to the two phrases, I performed two-speaker playback experiments. When the same numbers of phrases were presented per unit time, females preferred the song with trills to that without trills. This result may reflect female preference for songs with greater sound density. In subsequent playback experiments, I equalized the total sound duration per unit time (duty cycle) in songs with and without trills. The numbers of females that preferred songs with and without trills did not differ significantly. This suggests that trills can attract females like chirps do, even though the two sounds have different components.     

Assortative preferences and discrimination by females against hybrid male song in the grasshoppers Chorthippus brunneus and Chorthippus jacobsi (Orthoptera: Acrididae)

The grasshoppers Chorthippus brunneus and Chorthippus jacobsi are highly differentiated for male mating signals, and form a mosaic hybrid zone in northern Spain. At some sites within this zone, many hybrids are observed. At others, few hybrids are observed. Such bimodal sites may reflect recent contacts between parental genotypes, or local variation in levels of assortative mating or selection against hybrids. Playback of 12 parental and F1 male songs to 296 parental and hybrid females revealed positive assortative preferences in C. brunneus and C. jacobsi females, supporting a direct role of male mating signals in female choice. However, all female genotypic classes showed reduced responsiveness to F1 male songs. Such sexual selection against hybrids is consistent with the narrow cline width observed in the field for song characters relative to morphology. These results have implications for the genetic structure of the hybrid zone and for models of speciation by reinforcement.     

Host preferences in a phonotactic parasitoid of field crickets: the relative importance of host song characters

Abstract.  1. Predators, including insect parasitoids, often eavesdrop on prey signals, and as a result, predation can have important effects on the evolution of prey signalling behaviour.
2. The phonotactic parasitoid fly, Ormia ochracea , uses the calling songs of male field crickets to locate their field crickets hosts. In the western USA, this fly parasitises the variable field cricket, Gryllus lineaticeps . Previous work with one fly population suggested that female flies, like female field crickets, preferentially orient to male songs with higher chirp rates and longer chirp durations, although a limited range of male song types was used in this previous study. The current study, with a different fly population, used field-based, two-speaker choice tests to examine: (1) the effect of male chirp rate and chirp duration on fly attraction, using a natural range of song types; and (2) the relative importance of these song types in host selection by the flies.
3. Three lines of evidence suggested that chirp rate is more important than chirp duration in host selection. (a) The flies consistently preferred higher chirp rates but only sometimes preferred longer chirp durations. (b) The flies consistently preferred higher chirp rate/shorter chirp duration songs to lower chirp rate/longer chirp duration songs. (c) Preferences for longer chirp durations could be eliminated by increasing the amplitude of the less attractive song type, while preferences for higher chirp rates could only sometimes be eliminated by increasing the amplitude of the less attractive song type.
4. Fly predation may favour lower chirp rates and shorter chirp durations in G. lineaticeps , and may have resulted in stronger selection on chirp rate than on chirp duration.