Vibrations in the orbwebs of Nuctenea sclopetaria (Araneidae)

Summary Orbweaving spiders obtain much information from web-borne vibrations. One of their major problems is to distinguish relevant vibrations, particularly prey-produced signals, from irrelevant vibrations, particularly wind noise. This problem was studied by measuring the spectral characteristics of prey-and wind-induced web vibrations, and the spider's responsiveness to sinusoidal stimuli. In wind-generated vibration (Fig. 1) signal amplitude is large below 10 Hz and decreases by 30 to 40 dB/decade above the lowest resonance frequency for each type of web vibration. The lowest resonance for transverse vibration is 3 Hz with a 200 mg spider, and 10 Hz for longitudinal and lateral vibration. Below 10 Hz transverse vibration is much larger than the other two types, but this difference decreases at higher frequencies. Most of the amplitude in prey signals is below 50 Hz (Fig. 2), except for buzzing insects such as flies, bees and mosquitoes, whose high wing beats generate significant motion above 100 Hz. The spider's response threshold for sinusoidal stimuli in the prey-catching region of the web (Fig. 3) declines at 20 dB/decade over the range that could be measured (10 to 3,200 Hz for longitudinal, 18 to 560 Hz for transverse and 18 to 320 Hz for lateral vibration). The threshold curve for longitudinal vibration is 10 to 20 dB lower than for the other two types and has a minimum of 7 m RMS near 300 Hz. This difference appears to have a mechanical rather than a sensory basis, due to the better transmission of longitudinal vibration through the web (Fig. 4A). If the effect of the spider on web vibration is also considered, the resulting adjusted threshold curve (Fig. 4B: data available for longitudinal vibration only), shows a continuous decline of approximately 30 dB/decade. A possible mechanism for recognizing prey signals based on spectral information is discussed.     

A comparison of sound propagation and song frequency in temperate marsh and grassland habitats

Summary Attenuation of pure tones was measured in marsh and grassland habitat. At surface level, in grassland, the ground effect strongly attenuated frequencies below 2.0 kHz (Fig. 2). The ground effect was reduced by increasing source-receiver elevation. In marsh habitat the ground effect did not occur, and low frequencies were optimal for sound propagation (Fig. 3). As predicted from sound propagation tests, analysis of recorded songs of seven grassland and six marsh species indicated that minimum and emphasized frequency were significantly lower in songs of marsh birds. Maximum frequency did not differ between habitats (Table 4). Buzzing songs of yellow-headed blackbirds were broadcast and re-recorded at 1 and 50 m from the speaker in marsh and grassland habitats. Low frequency components attenuated more rapidly than higher frequency components in grassland, and the reverse occurred in marsh. Results suggested that the ground effect restricts the use of low frequencies by grassland birds. This evidence is consistent with the sound window hypothesis of Morton.     

Roles of interplant movement,acoustic communication,and phototaxis in mate-location behavior of the leafhopper Graminella nigrifrons

Summary Male Graminella nigrifrons leafhoppers (Cicadellidae: Homoptera) employ a call-fly strategy to find virgin females on oat host plants. Males observed in isolation during daylight hours exhibit a high rate of interplant movement, calling from the lower canopy on each plant visited. Virgin and mated females exhibit little interplant movement. They differ from one another in that virgin females perch on the upper half of plants, whereas mated females perch on the lower half of plants. The positioning of females in the plant canopy is influenced by light. Unlike mated females, virgin females respond to male calls by emitting their own acoustic signals. When virgin females are present on plants visited by males, interplant movement of males ceases, and a localized-upward search of the female bearing plant ensues. Male search is influenced by light. Regardless of whether virgin females were confined to the upper or lower portion of plants, direction of male search was towards a light source used to illuminate above or below the plant canopy. These findings suggest that interplant movement by males and sedentary behavior by females prior to mate recognition and their use of acoustic and phototactic sensory modalities after mate recognition represent previously unrecognized adaptations to problems associated with the use of vibrational signals on plants. Offprint requests to: R.E. Hunt at the current address     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

High-frequency sound transmission in natural habitats: implications for the evolution of insect acoustic communication

Summary Transmission and reception of high-frequency sound in the natural environment of bushcrickets (Tettigonia viridissima L.) was studied using the activity of an identified neuron in the insect's auditory pathway as a biological microphone. Different positions of the receiver within the habitat were simulated by systematic variation of the distance from a loudspeaker and the height above the ground. Attenuation and filtering properties of the habitat were investigated with pure-tone frequencies between 5 and 40 kHz. Sound attenuation in excess of the attenuation due to geometrical spreading alone increased with increasing frequency, distance between sender and receiver, and decreasing height within the vegetation (Figs. 2–4). The data also confirm the existence of two kinds of excess attenuation. The amount of amplitude fluctuations in the sound signals was investigated by analysing the variability of the neuronal responses at a given receiver position. Variability increased with decreasing bandwidth of a noise signal at some distance from the loadspeaker. The variability in the responses to pure tones increased with both increasing frequency and distance from the source (Fig. 7). In the selected habitat, the temporal pattern of the natural calling song of male T. viridissima was very reliably reflected in the activity of the recorded neuron up to a distance of 30 m at the top of the vegetation, and 15–20 m near ground level (Figs. 5, 8). The maximum hearing distance in response to the calling song was about 40 m. Environmental constraints on long-range acoustic communication in the habitat are discussed in relation to possible adaptations of both the signal structure and the behavior of the insects.Abbreviations excessive attenuation EA - sound pressure level SPL Offprint requests to: H. Romer     

Temporal patterning of within-song type and between-song type variation in song repertoires

In many songbird species, males sing a repertoire of distinct song types. Song sparrows typically are described as having repertoires of about a dozen song types, but these song types are themselves quite variable and some songs are produced that appear intermediate between two types. In this study we quantify the similarity between successive songs produced by song sparrows in order to determine if differences between song types are emphasized or deemphasized in bouts of continuous singing. In spite of the high degree of variation within song types and similarity between song types observed in this species, we show that transitions from one song type to the next are distinct as compared to transitions within sequences of the same type (Figs. 4 and 5). Variation does not accumulate across sequences of the same song type, and the average variation observed within a continuous sequence of the same song type is significantly less than is predicted from the total variation recorded for that type across many different bouts (Fig. 6). These results support the view that song sparrows include two levels of variation in their singing: differences between song types as is commonly observed in species with song repertoires, and an independent level of variation observed for songs of the same type.     

Multimodal signals increase active space of communication by wolf spiders in a complex litter environment

Multimodal signals may compensate for environmental constraints on communication, as signals in different modalities vary in efficacy. We examined the influence of complex microhabitats on transmission of vibratory and visual signals of courting male Schizocosa ocreata wolf spiders (Araneae: Lycosidae) with laser Doppler vibrometry (LDV) and behavioral observations in lab and field. We measured maximum potential detection distance of visual and vibratory signals by females in laboratory mesocosms, recorded vibration signal attenuation on different substrates, and estimated transmission distances for male vibration signals in the field. We also determined effective line-of-sight visual detection distances in the field with laser distance measures. Together, these data were used to estimate the potential and effective active space of multimodal signals. LDV measures show leaves are highly conductive substrates for wolf spider vibratory signals compared to others (soil, wood, rock). For both visual and vibratory modes, lab estimates of maximum potential distance for signal transmission and detection (behavior studies) exceeded estimates of effective active space (signal attenuation, “vanishing point,” and “line-of-sight” measures). Field estimates of transmission distance for signal modes overlap, such that in close range (<20 cm), vibratory signals are more likely to be detected, while farther away, visual signals are more likely to be seen. These findings thus support current hypotheses regarding how multimodal communication might extend the range of overall signal active space or compensate for environmental constraints.     

Blue-winged warblers (Vermivora pinus) “recognize” dialects in type II but not type I songs

Summary Male blue-winged warblers (Vermivora pinus) sing two distinctly different song types (I and II). Type II songs from eastern New York and western Massachusetts, USA, are different (Fig. 1), and during playback experiments males respond stronger to local than to foreign songs (Fig. 2). Type I songs are more stereotyped geographically (Fig. 1), and males do not differentiate between local and foreign Type I songs (Fig. 3). Differing degrees of intra- and inter-sexual selection on these two forms may have promoted variability in Type II songs and stereotypy in Type I songs among locations.     

Dominions and desert clickers (Orthoptera:Acrididae): influences of resources and male signaling on female settlement patterns

Summary Previous field work on the grasshopper Ligurotettix coquilletti revealed that females were not evenly distributed among male mating territories, Larrea tridentata (creosote) bushes, but were clustered at particular locations. These sites generally harbored several signaling males simultaneously and also possessed foliage preferred by the insects as a food source, this preference being based on the relative concentrations of various extra-foliar compounds. The clustering of females, therefore, could result from a preference for specific bushes because of the resources (i.e., food) available there and/or an orientation to groups of males per se. Here, we present the results of 3 field experiments in which we controlled the spatial distribution and intensity of male signals using a computer-operated system of loudspeakers and monitored the movement of individually marked females released in the study area. When male song was identical at high and low quality territories (all having single loudspeakers), females still aggregated at the high quality sites, indicating that variability in host plant quality alone may be sufficient to promote a skewed distribution of females. Among high quality territories, females did not discriminate between sites with one versus three loudspeakers (all broadcasting the same signal), but displayed a strong preference for sites (all having single loudspeakers) with a high intensity signal over a low intensity one. Field measurements showed that the songs of grouped males were more intense than those of lone males, implying that the signaling of grouped males may have enhanced the settlement of females at the bushes harboring male groups above and beyond that influenced by territory quality alone. We conclude that female attraction to host plants is influenced primarily by male signaling, whereas their subsequent retention is more dependent on territory quality. An experiment on male settlement failed to show an aggregative tendency, suggesting that male groups form through the passive accumulation of individuals at high quality sites.     

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

Geographic variation in songs of the Bewick's wren: a search for correlations with avifaunal complexity

Summary Song characteristics of the Bewick's wren (Thryomanes bewickii) are compared from nine localities in the western United States. Character shifts, i.e., a difference in means, are evident for all song characters: Arizona and Colorado songs are especially short and long, respectively (Figs. 1, 2); songs of insular (Santa Cruz Is.) and nearby mainland populations in California are very dissimilar (Table 1); excluding the insular population, the frequency range of song phrases is positively correlated with latitude (Fig. 3). Variance shifts, i.e., a difference in repertoire size or Coefficients of Variation (CV's) of measured song characters, are also present; most notably, Arizona males have exceptionally stereotyped songs, with small song phrase repertoires (Table 2) and low CV's. Population densities and/or habitat structure undoubtedly influence signal design, but correlations reported here suggest that the avifaunal complexity and the corresponding vocal milieu should also be examined rigorously as possibly important influences.     

Sensory ecology of a semi-aquatic spider (Dolomedes triton)

Summary Preferred ambush sites of fishing spiders Dolomedes triton (Pisauridae) were investigated, primarily with respect to aquatic vegetation and wave action, at five locations differing in these variables along the shore of Dow Lake (near Athens, Ohio). The linear density of spiders correlated positively with the presence of aquatic vegetation. D. triton most often stood at a distance from shore proportional to the width of the zone of floating and emergent vegetation (Fig. 2). Usually they positioned their bodies in the horizontal plane on floating living or dead plant parts. Spider density was negatively correlated with the amplitude of wind-generated waves (Fig. 4). The maximum energy of these waves measured on breezy days was at a frequency of 1.4 Hz, with no components exceeding 10 Hz (Fig. 5). Spiders could discriminate high-amplitude (up to 40 mm) wind waves from prey waves, probably since the latter have substantially higher frequencies (Figs. 3, 5).     

Inhibition of glycosidases by Lepidoptera; roles in the insects and leads to novel compounds?

Summary Glycosidase inhibitors are widespread in plants and can be sequestered by Lepidoptera, for which they can presumably serve as defences by making the insects indigestible to a range of potential predators. As a result of this study of eight British species of moth and butterfly it was found that glycosidase inhibitors in the insects could then be detected in the larval food plants which were not previously known to contain them; however, some were only detectable in the plants after concentration. In some cases the inhibition of specific glycosidases by Lepidoptera was detected even though the insects had not apparently acquired them from their food plants. Inhibition of-N-acetylglucosaminidase was observed in most of the adult Lepidoptera analysed but further work is required to identify the inhibitors, though they are likely to be nitrogen-containing compounds. Weak anti-HIV activity was also observed in the glycosidase-inhibiting fractions ofAcherontia atropos and the plantUrtica dioica.     

Acoustic communication in phaneropterid bushcrickets: species-specific delay of female stridulatory response and matching male sensory time window

Summary The stridulatory sounds and movements produced by the females of various bushcricket species (Tettigonoidea: Phaneropteridae: Barbitistini) are compared with those of the males. Behavioral experiments are carried out to determine the significance of the female sounds in acoustic communication. Selection factors involved in the evolution of female stridulation are discussed. The morphological apparatus for sound production has evolved independently in males and females. Whereas males rub a toothed file on the underside of the left wing over the inner edge of the right wing, the plectrum, the females stridulate by rubbing a thickened vein on the underside of the left wing over modified spines on the upper surface of the right wing (Fig. 2). Similarly, the movements responsible for sound production are not homologous in males and females. In the male the audible closing movement is always preceded by wing opening, whereas the female in general initiates the closing movement when the wings are in the resting slightly opened position, and abruptly produces complete closure (Fig. 3, 4 and 5). The female responds to male singing by emitting one to several highly damped sound pulses each lasting less than one ms. The interval between the song of a conspecific male and this response is a very precise species-specific characteristic (Fig. 7). In species with male songs that are complicated in structure or continuous, the females respond only at specific time-after patticular markers in the song of the male. The time interval between male song and female response is an important criterion by which the male identifies conspecific female song (Fig. 8). Because the response delay of the female and corresponding neuronal time window in the male are distinct they may be important in species discrimination.     

Echolocation in two very small bats from Thailand Craseonycteris thonglongyai and Myotis siligorensis

Summary The echolocation and hunting behavior of two very small bats, Craseonycteris thonglongyai (Hill) and Myotis siligorensis (Horsfield), from Thailand, were investigated using multiflash photographs, video, and high-speed tape recordings with a microphone array that allowed determination of distance and direction to the bats. C. thonglongyai is the world's smallest mammal and M. siligorensis is only slightly larger. Both bats hunted insects in open areas. The search signals of C. thonglongyai were 3.5 ms long multiharmonic constant frequency (CF) signals with a prominent second harmonic at 73 kHz repeated at around 22 Hz. The band width (BW) of the short terminal frequency modulated (FM) sweep increased during the very short approach phase. In the final buzz the CF component disappeared, the duration decreased to 0.2 ms, and the repetition rate increased to 215 Hz (Figs. 2, 3, 4). There was no drop in frequency in the buzz. The video recordings of C. thonglongyai indicated that it seizes insects directly with the mouth (Fig. 1). M. siligorensis produced 5.4 ms long CF search signals at 66 kHz. The repetition rate was around 13 Hz. In the approach phase an initial broad band FM sweep was added. The buzz consisted of two phases, buzz I and buzz II. Buzz 11 was characterized by short cry durations (around 0.3 ms), a constant high repetition rate (185 Hz), a distinct drop in frequency, and a prominent second harmonic (Figs. 5, 6, 7). The drop in frequency, apparently typical of vespertilionid bats, has been explained by physiological limitations in sound production. However, C. thonglongyai produced very short signals at very high repetition rates without any frequency drop. The drop may be of adaptive value since it enables M. siligorensis to produce very short signals with high sweep rates. The drop moves the pronounced second harmonic into the frequency range of most interest to the bat (Fig. 7D). The sweep rate in this frequency range may now increase to twice the maximum rate that the vocal cords can produce directly. C. thonglongyai and M. siligorensis belong to different superfamilies, Emballonuroidea and Vespertilionoidea, respectively. In spite of their phylogenetic distance they produce strikingly similar search signals of narrow BW around 70 kHz with high source levels (100–115 dB peSPL peak equivalent sound pressure level). We argue that the signal resemblance is due to the similarity in size and hunting behavior of the two bats both hunting insects in open areas. High frequencies are heavily attenuated in air, but because of their small size the bats are restricted to hunting small insects which only reflect echoes at high frequencies. Thus, the emitted frequency is probably the lowest possible given the prey size. Hence, the two bats can only maximize the range of their sonar by decreasing the BW and emitting high intensities. Correspondence to: A. Surlykke     

Infrasonic calls of the Asian elephant (Elephas maximus)

Summary Calls at frequencies below the range of human hearing were recorded from two groups of captive Asian elephants (Elephas maximus). Most of the calls ranged in frequency from 14 to 24 Hz, with durations of 10–15 s (Fig. 1). With the nearest elephant 5 m from the microphone, sound pressure levels were 85 to 90 dB (re 20 Pa). These calls occurred in a variety of circumstances. Elephants are the first terrestrial mammals reported to produce infrasound. These calls may be important in the coordination of behavior in thick vegetation or among separated groups of elephants.     

Dragon’s blood secretion and its ecological significance

Dragon’s blood is the name given to a red exudate produced by some plant species belonging to the genera Daemonorops, Dracaena, Croton and Pterocarpus. These are endemic to various parts of the globe. It is classified as a resin or latex depending on its mode of secretion and its chemical composition, which is species specific. This red substance functions in defence and is produced (a) constitutively and stored in preformed anatomical structures, or (b) by induction in response to traumatic events, such as mechanical injury, pathogen attack or invasion by insects. Apart from its defensive role in plants, dragon’s blood is also a valuable natural resource renowned since antiquity for its diverse medicinal properties and uses in art. Despite the great importance of dragon’s blood, our knowledge of the biological basis for its secretion is still incomplete. This review summarizes recent advances in the study of the anatomical basis for its secretion, and discusses its classification and ecological function. Bringing some clarity to these issues may also help in the commercial sourcing of dragon’s blood.     

Performance evaluation of circulating fluidized bed incineration of municipal solid waste by multivariate outlier detection in China

This first nationwide survey was conducted to evaluate the overall performance of the circulating fluidized bed (CFB) incineration of municipal solid waste (MSW) in 2014-2015 in China. Total 23 CFB incineration power plants were evaluated. The data for monthly average flue gas emission of particles, CO, NO _x , SO₂ and HCl were collected over 12 consecutive months. The data were analyzed to assess the overall performance of CFB incineration by applying the Mahalanobis distance as a multivariate outlier detection method. Although the flue gas emission parameters had met the Chinese national emission standards, there were 11 total outliers (abnormal behavior) detected in 6 out of 23 CFB incineration power plants from the perspective of the MSWincineration performance. The results demonstrate that it is more important for a better performance of CFBs to reduce the frequencies of the MSW load changes, rather than the magnitudes of the MSW load changes, particularly reducing the frequencies in the range of 10% and more of the load changes, under the same and stable conditions. Furthermore, the overloading occurs more often than the underloading during the operation of the CFB incineration power plants in China. The frequent overloading is 0% to 30% of the designed capacity. To achieve the stable performance of CFBs in practice, an appropriately designed MSW storage capacity is suggested to build in a plant to buffer and reduce the frequency of the load changes.

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Survival of first instar larvae ofDanaus plexippus (Lepidoptera: Danainae) in relation to cardiac glycoside and latex content ofAsclepias humistrata (Asclepiadaceae)

Summary Our paper addresses field survivorship of first instar monarch butterfly larvae (Danaus plexippus L., Lep.: Danainae) in relation to the dual cardenolide and latex chemical defenses of the sand hill milkweed plant,Asclepias humistrata (Asclepiadaceae) growing naturally in north central Florida. Survival of first instar larvae in the field was 11.5% in the first experiment (15–20 April 1990), and dropped to 3.4% in the second experiment (20–30 April). About 30% of the larvae were found glued to the leaf surface by the milkweed latex. Predator exclusion of non-flying inverte-brates by applying tanglefoot to the plant stems suggested that the balance of the mortality was due to volant inverte-brates, or to falling and/or moving off the plants. Regression analyses to isolate some of the other variables affecting survivorship indicated that first instar mortality was correlated with (1) increasing cardiac glycoside concentration of the leaves, (2) increasing age of the plants, and (3) the temporal increase in concentration of cardiac glycosides in the leaves. The study also provided confirmatory data of previous studies that wild monarch females tend to oviposit onA. humistrata plants containing intermediate concentrations of cardiac glycosides. Cardiac glycoside concentration in the leaves was not correlated with that in the latex. The concentration of cardenolide in the latex is extremely high, constituting an average of 1.2 and 9.5% of the mass of the wet and dry latex, respectively. The data suggest that an increase in water content of the latex is compensated for by an influx of cardenolide with the result that the cardenolide concentration remains constant in the latex systems of plants that are growing naturally. We also observed first instar larvae taking their first bite of milkweed leaves in the field. In addition to confirming other workers findings that monarch larvae possess elaborate sabotaging behaviour of the milkweed's latex system, we discovered that several larvae on their first bite involuntarily imbided a small globule of latex and instantly became cataleptic. This catalepsis, lasting up to 10 min, may have been in response to the high concentration of cardenolide present in the latex ofA. humistrata, more than 10 times that in the leaves. The results of the present study suggest that more attention should be directed to plant chemical defenses upon initial attack by first instar insect larvae, rather than attempting correlations of plant chemistry with older larvae that have already passed the early instar gauntlet. The first bite of neonate insects may be the most critical moment for coping with the chemical defenses of many plants and may play a much more important role in the evolution of insect herbivory than has previously been recognized.     

The response of female Mountain White-crowned Sparrows to songs from their natal dialect and an alien dialect

Summary Female Mountain White-crowned Sparrows from Colorado were tested in a laboratory playback experiment to determine their response to male songs from their own natal dialect and to those from an alien dialect of the same subspecies, Zonotrichia leucophrys oriantha (Fig. 1). Observations were made on locomotor activity and copulation postures produced during each experimental test session.The subjects gave copulation displays almost exclusively when hearing their natal dialect and new or no displays when hearing the alien dialect (Fig. 2). The birds also exhibited significantly greater locomotor activity when hearing the natal dialect than they did when hearing the alien dialect (Fig. 3).We conclude that females are sensitized by their early song learning experience to be responsive to songs from their natal dialect and virtually sexually unresponsive to songs from an alien dialect. We speculate that the copulation posture given in the experiment implies that females in natural populations would preferably only mate assortatively with males from their natal dialect region.