The effectiveness of katydid (<Emphasis Type="Italic">Neoconocephalus ensiger</Emphasis>) song cessation as antipredator defence against the gleaning bat <Emphasis Type="Italic">Myotis septentrionalis</Emphasis>

Many nocturnal katydids (Orthoptera: Tettigoniidae) produce intense calling songs, and some bat species use these songs to detect and locate prey. One Nearctic katydid species, Neoconocephalus ensiger, ceases or pauses singing in response to bat echolocation calls. We tested the hypothesis that song cessation is an effective defence against gleaning bats (i.e., bats that take prey from surfaces). We observed Myotis septentrionalis, a sympatric bat species that uses prey-generated sounds when gleaning, attack and feed on singing N. ensiger in an outdoor flight room. These bats demonstrated a preference for the calling song of N. ensiger over a novel cricket calling song when they were broadcast from a speaker in the flight room. Bats attacked speakers broadcasting N. ensiger calling song as long as the song was continuous and aborted their attack if the sound stopped as they approached, regardless of whether a katydid was present as a physical target on the speaker. Echolocation calls were recorded during attacks and no significant differences were found between continuous and interrupted song approaches for four call parameters, suggesting that M. septentrionalis may not use echolocation to locate silent prey. Therefore, song cessation by katydids in response to ultrasound is an effective defence against gleaning bats.     

A dusk chorus effect in a nocturnal bird: support for mate and rival assessment functions

The dawn chorus is a striking feature of spring mornings and a characteristic behaviour of many bird species, particularly the passerines. Dawn singing has been considered a reliable signal of male quality for mate and rival assessment. Singing is presumed to be relatively costly at dawn both because air temperatures are relatively low and because birds have not fed overnight. Models of optimal daily routine predict the existence of a “dusk chorus” in nocturnal birds, although this prediction has received little empirical attention. Nocturnal birds at dusk may be energy-limited because of a lack of daytime feeding, and singing at dusk may thus ensure signal reliability. Here, we used an observational and experimental approach to study vocal behaviour at dusk and dawn in a nocturnal raptor, the little owl Athene noctua. We assess whether male little owls adjust their vocal behaviour according to feeding stage (i.e. period of the night), ambient air temperature and territorial context (i.e. spontaneous calling behaviour vs elicited calling by intrusion). Across different temperatures, we find that both spontaneous vocal activity and inter-individual variability in call duration increased at dusk, clearly indicating a dusk chorus phenomenon. Results from playback presentations suggest that food, rather than air temperature, is likely to be more constraining at dusk. We discuss how comparing dusk and dawn choruses in nocturnal and diurnal species can provide insights into both mechanistic and functional aspects of signalling behaviour.     

An attractive male trait and aggressiveness are negatively correlated in wild field crickets, but uncorrelated in lab-reared crickets

It is generally thought that attractive male traits are positively correlated with dominance (aggressiveness). However, growing evidence suggests that this is not necessarily the case. We investigated whether calling song, a male mating display used by females to evaluate potential mates in a field cricket (Gryllus integer), is correlated with aggressiveness. In this species, females prefer males with longer durations of singing time. We measured singing time by measuring song over three continuous days using a custom-designed audio-monitor and assessed aggressiveness by pairing males in agonistic interactions. Our results showed that for males caught in the field, the percentage of time spent singing was negatively correlated with aggressiveness. However, in males that were raised in the laboratory, the percentage of time spent singing was not correlated with aggressiveness. Since calling duration is an attractive male trait in these crickets, our data show that the attractiveness of males can be either negatively related to aggressiveness (field) or unrelated to aggressiveness (lab). Neither of these results fit the traditional view that preferred male mating cues should be positively correlated with dominance (aggressiveness). These results also suggest that measurements of aggressiveness made in the lab do not necessarily correspond with aggressiveness in the field.     

Carbohydrate and calling: Depletion of muscle glycogen and the chorusing dynamics of the neotropical treefrog Hyla microcephala

Chorusing males of the neotropical treefrog Hyla microcephala call in distinct bouts punctuated by periods of silence, a pattern known as unison bout singing. Schwartz (1991) previously tested and refuted the hypotheses that males periodically stop calling either because of a female preference for males that call cyclically, or because high ambient noise levels inhibit vocal activity. Males of H. microcephala are vocally responsive to the calls of other males, and during calling bouts their rate of note production can exceed 10,000 per hour. In natural choruses females preferentially pair with males that call at the higher rates. Because females can pair with males over many hours, males may stop calling periodically to save energy so they can continue to call for the entire period that females are available. We directly tested this energy conservation hypothesis by collecting samples of males early in the evening just after chorusing commenced and later when chorusing had ended for the night. Trunk muscles (internal and external oblique), which are responsible for the airflow associated with note production, were dissected, frozen, and their glycogen content measured. Data on calling behavior were obtained for late-evening samples. Individual calling behavior was not correlated with a males final glycogen level. In addition, many males ended their calling before glycogen reserves were exhausted, indicating that factors other than energy can determine when males finally stop chorusing, However, the biochemical assays supported the energy conservation hypothesis. Unless chorusing was punctuated by pauses, most males would have been unable to sustain high rates of calling for an entire evening without exhausting glycogen reserves in their trunk muscles. Because the time females pair with males is probably unpredictable to males, the ability to call for long periods may improve a males chances of mating.     

Bat predation and the evolution of leks in acoustic moths

Theories of lek evolution generally invoke enhanced mating success experienced by males signalling in aggregations. Reduced predation has also been acknowledged as a potential factor driving lek formation, but its role is more ambiguous. Although lekking is a complex behaviour, few empirical studies have investigated the role of both claims. We studied the potential pressures imposed by mating success and predation in an acoustic moth, Achroia grisella, in which males gather in leks and broadcast a calling song attractive to females. We exploited the ability to manipulate the distribution of singing males in laboratory arenas to create different-sized leks and tested female preferences for these aggregations. Because A. grisella are vulnerable to predation by bats while in flight and on the substrate, we also tested the responses of a potential predator, Rhinolophus ferrumequinum, a bat species that feeds on moths, to the experimental leks. We found that the per capita attractiveness of A. grisella males to females rose with increasing lek size. R. ferrumequinum also oriented toward experimental A. grisella leks, but this attraction did not increase at larger leks. Thus, a male’s per capita exposure to predation risk declined as more moths joined the lek. A. grisella males appear to benefit from advertising in larger leks in terms of both increased mate attraction and reduced predation risk. Our results support the idea that multiple factors operating simultaneously may maintain lekking behaviour.     

Reproductive strategy and singing activity: blue tit and great tit compared

The costs and benefits of bird song are likely to vary among species, and different singing patterns may reflect differences in reproductive strategies. We compared temporal patterns of singing activity in two songbird species, the blue tit (Cyanistes caeruleus) and the great tit (Parus major). The two species live side by side year round, and they have similar breeding ecology and similar rates of extra-pair paternity. However, they differ in two aspects of reproductive strategy that may have an influence on song output: blue tits are facultatively polygynous and have a fairly short breeding season with almost no second broods, whereas great tits are socially monogamous but more commonly raise second broods. We found that great tit males continued singing at high levels during the egg-laying and incubation periods, while monogamously paired blue tit males strongly reduced singing activity after the first days of egg-laying by their female. Since males of both species sang much more intensely shortly before sunrise than after sunrise, at midday or in the evening, this difference was most conspicuous at dawn. No differences in singing activity were found within species when testing for male age. We suggest that in contrast to blue tits, great tit males continued singing after egg-laying to defend the territory and to encourage the female for a possible second brood.     

Aggressive thresholds in Dendropsophus ebraccatus: habituation and sensitization to different call types

Males in many chorusing anuran species use aggressive calls during defense of calling spaces from other males. The minimal intensity of another male’s vocalizations that elicits an aggressive call response has been termed the aggressive threshold. Previous studies of aggressive thresholds have shown that they are plastic: males habituated (increased their aggressive thresholds) in response to repeated presentation of stimuli above initial threshold levels. Habituation likely contributes to the stable chorus structure of these species, in which aggressive calling is rare compared to advertisement calls. I have observed high levels of aggressive calling in the treefrog Dendropsophus ebraccatus, suggesting that males of this species do not habituate. In this study, I investigated the plasticity of aggressive thresholds in D. ebraccatus. I measured the aggressive thresholds of males before and after suprathreshold stimulation by both advertisement and aggressive calls. I found that the different call types had different effects: males habituated to advertisement calls but lowered their aggressive thresholds in response to aggressive calls. I consider the latter response to be an example of sensitization, a behavior that has been documented infrequently in vocalizing anurans. Sensitization is a plausible mechanism responsible for the high levels of aggressive calling observed in this species. Given the high costs of aggressive calling, however, it is unclear why a mechanism that increases aggressive call output would be maintained.     

Latency to resume calling after disturbance in the field cricket,<Emphasis Type="Italic"> Teleogryllus oceanicus</Emphasis>, corresponds to population-level differences in parasitism risk

A possible parasitoid-evasion behavioral adaptation is examined in male field crickets, Teleogryllus oceanicus, from three Hawaiian islands where parasitoid prevalence varies naturally among islands. Ormia ochracea, the parasitoid fly that parasitizes T. oceanicus on these islands, uses male calling song to locate its hosts. We used laboratory-reared males from three Hawaiian islands to determine if there are population differences in the time it takes for calling males to resume calling after a standardized disturbance. Males follow the expected pattern; males from the island with the greatest risk of parasitism have the longest latency to resume calling, and males from the island with the least risk of parasitism have the shortest latency to resume calling. Results are discussed in the context of behavioral adaptations to differing parasitism levels, and trade-offs between natural and sexual selection.Communicated by D. Gwynne     

Alarm calling behavior of the thirteen-lined ground squirrel,Spermophilus tridecemlineatus

Summary Alarm calling in a population of thirteenlined ground squirrels, Spermophilus tridecemlineatus, was studied over a three-year period. Data on ground squirrel reactions to human and canine approaches and to the approach or presence of avian predators were used to quantify alarm calling behavior.The results support the hypothesis that alarm calling in this species functions to warn genetic relatives. Human and canine approach-elicited calls were most frequently given by mothers and their recently emerged young; adult males and nonparous females rarely called. The onset of maternal calling coincided with the aboveground appearance of a mother's own litter, and both juvenile and maternal calling were terminated at approximately three weeks post-emergence. Alarm calls were rarely emitted during encounters with avian predators.Alarm calling behavior in S. tridecemlineatus thus appeared to be dependent upon the presence of newly emerged juvenile relatives. To investigate whether the population structure of S. tridecemlineatus was perhaps incompatible with the evolution of alarm calling directed toward adult relatives, the distance between the home ranges of adult relatives and the distance over which the alarm vocalization is audible to ground squirrels were measured. The results revealed that females were likely to have adult relatives relatives residing within audible range of the call.     

The effect of mating on female sexual behavior of Teleogryllus commodus Walker

Summary Virgin Teleogryllus commodus females show rhythmic locomotor activity in light-dark (LD) and constant light (LL) conditions. A taped 3-h calling song stimulates them into further walking, and they lay few eggs. Successful mating suppresses or reduces circadian-controlled walking, but in LL-conditions the free-run still persists. Mated females no longer respond to the 3-h calling song. The number of eggs laid within 24–72 h after insemination increases, and the three effects of mating occur independently of one another. The possible physiological basis of these changes and their ecological and behavioral significance under field conditions are discussed.     

Divergence in female calling song discrimination between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae)

Two species of closely related wood cricket, Gryllus fultoni (Orthoptera: Gryllidae) and Gryllus vernalis, occur together in some parts of the eastern United States and have a similar calling song structure, consisting of three-pulse chirps. A previous study revealed that chirp rate in G. fultoni was highest (greatest difference vis-à-vis chirp rate in G. vernalis) in sympatric populations, intermediate in near allopatric populations that were located close to the sympatric zone, and lowest in allopatric populations. A similar trend was observed in pulse rate, but the mean values of this trait showed much more convergence than chirp rate at the low end of the range of calling temperatures. In this study, we investigated the song discrimination of females from sympatric and allopatric populations of G. fultoni at about 23°C, which is near the middle of the normal range of calling temperatures. We used both single-stimulus and two-stimulus playback experiments to learn if geographical differences in song preferences paralleled those in calling songs. Stimuli presented were representative of calling songs in three classes of G. fultoni populations (sympatric, near allopatric, and far allopatric), a calling song of G. vernalis, and three calling songs with parameter values that were intermediate with respect to those of the songs of far allopatric G. fultoni and G. vernalis. In the single-stimulus playbacks, females of all G. fultoni populations responded poorly if at all to the heterospecific stimulus. Females of sympatric and near allopatric populations responded poorly to all intermediate stimuli, but females of far allopatric populations frequently responded to these sounds. In the two-stimulus playbacks, females of sympatric and near allopatric populations generally discriminated against intermediate and heterospecific stimuli. However, females of far allopatric populations often did not discriminate against intermediate stimuli, whose characteristics resembled the calling songs of G. vernalis. The divergent pattern of female phonotactic discrimination between sympatric and far allopatric populations was thus generally congruent with the pattern of divergence in chirp and pulse rates and would be expected to significantly reduce heterospecific mating in sympatry. These geographical patterns of female song discrimination and male calling songs conform to a commonly used definition of reproductive character displacement.     

Interactions between singing Hawaiian humpback whales and conspecifics nearby

Summary Interactions of singing humpback whales, Megaptera novaeangliae, with conspecifics nearly were studied during the breeding season off the west coast of Maui, Hawaii. On 35 occasions singing humpbacks were followed by boats (Table 1). The movement patterns of these singing whales and other conspecifics nearby were recorded by observers on land using a theodolite.Thirteen of 35 singers stopped singing and joined with nonsinging whales either simultaneously or within a few minutes after ceasing to sing. Another 15 also stopped singing while under observation and were not seen to join with another whale, but all singing whales that joined with other whales stopped singing. Singing whales often pursue nonsinging whales, while nonsinging whales usually turn away from singers (Figs. 4, 5).When a singer joined with a female and calf unaccompanied by another adult, behavior tentatively associated with courtship and mating was observed (Fig. 7). Such behavior also occurred during several interactions between singers and individuals of unknown sex. Aggressive behavior was observed during three interactions between singers and individuals of unknown sex (Fig. 4) and it predominated whenever more than one adult accompanied a cow and calf. During the other occasions when a singer joined another whale, we could not determine the nature of the interaction. Many times the singers and joiner would surface together only once and would then separate. However, on several occasions the singer and joiner would remain together for as long as we could follow them, up to 1.5 h.The roles of singer and joiner can be interchangeable. For instance, on two occasions a singer joined with a whale that either had been singing or started singing later in the day (Fig. 3). Furthermore, on several occasions, a nonsinging whale appeared to displace the singer. Individual singing humpbacks are not strictly territorial, although singers appear to avoid other singers.As the breeding season progressed, singers sang for longer periods of time (Fig. 2). In addition, the probability of a whale joining with the singer decreased by 42% from the first half of the observation period to the second half. Furthermore, this increase in duration of song bouts occurred during that section of the season when female reproductive activity as measured by rate of ovulation is reported to be decreasing in other areas.Our observations support the hypothesis that humpback song plays a reproductive role similar to that of bird song. Humpbacks sing only during the breeding season. If, as seems likely, most singing humpbacks are male, then singing humpbacks probably communicate their species, sex, location, readiness to mate with females, and readiness to engage in agonistic behavior with other whales.     

Choice of singing sites by male bushcrickets (Tettigonia viridissima) in relation to signal propagation

Summary The spatial dispersion of singing male bushcrickets (Tettigonia viridissima) in a marshland habitat was found to be significantly clumped. Males clustered in patches of taller vegetation, buth within such clusters, males were regularly spaced with a mean distance of 6 m between nearest-neighbours. Males occupied perches on plants that were, on average, 0.3 m above the top of surrounding vegetation. Physical and acoustical interactions between males were observed more frequently when males were singing from higher sites. Excess attenuation of the male song was found to increase with frequency but decreased markedly with increasing elevation of the singing male above the ground. The maximum detection range of the song, realized when the insect was singing > 1 m above the surrounding vegetation, was estimated as 60 m for the fundamental frequency (10 kHz), 38 m for the 1st harmonic (20 kHz) and 14 m for the second harmonic (30 kHz). By contrast, when males sang from the middle of dense reed beds, the estimated detection distance was only 8 m, 6 m and 4 m for each frequency band, respectively. Males could have increased the detection range of their songs almost three fold by singing from higher positions than those usually observed in the field. This suggests that there may be a cost of singing at higher elevations such as an increased risk of predation and/or increased aggression from neighbours. We suggest the spacing strategy adopted by males reflects a compromise between maximizing the range over which their songs can be detected and accurately localized by females and minimizing interference from competing males. Offprint requests to: T. Eiriksson     

Calling energetics of a neotropical treefrog,Hyla microcephala

Summary We investigated the calling energetics of Hyla microcephala, a neotropical treefrog with an unusually complex vocal repertoire. Males respond to the calls of other individuals by adding secondary click notes to their calls, thereby increasing the total number of notes given per minute. Rates of oxygen consumption of males calling in metabolic chambers were 0.41–2.80 ml O₂/(g·h), corresponding to calling rates of 205–6330 notes/h. Note rate explained 95% of the variance in meta-bolic rate; the effect of variation in body size and temperature was small. Data from playback experiments with males in the field showed that note rate increased as as the number of notes in a stimulus call increased, and this resulted in substantial increases in the cost of calling. Average metabolic rates for males in the field were about 1.7 ml O₂/(g·h), for a net cost of calling of about 20 J/h for an average-size male. However, estimated metabolic rates varied by more than 300% and were strongly influenced by the proximity and calling activity of other males in the chorus. Male H. microcephala appear to conserve energy by reducing calling rates when only a few males are active and increasing calling efforts only when vocal competition among males is intense.     

The adaptive significance of acoustic spacing in male bushcrickets Tettigonia viridissima: a perturbation experiment

Summary A perturbation experiment was carried out in which the spacing between singing male bushcrickets, Tettigonia viridissima, was artificially manipulated. The experiment entailed releasing virgin females into an area in which caged males were either spaced evenly or in which a proportion of the males were clumped. There was large variation among males in the proportion of time spent singing (range 39–91% of total singing time). Singing activity was correlated with male body weight, but was not influenced by male spacing. Out of a total of 108 females released during the experiment, the majority (N=90) moved from their release points onto the cage of one of the closest singing males. More distant signalers sometimes attracted females when one of the closest males did not sing, or sang very little, during the period in which females were moving. When clumped, males were less successful in attracting females than when regularly spaced within the experimental area. Therefore, within a homogeneous habitat in which females are randomly distributed, male mating success will be maximized when males space out as far as possible from their competitors. As predicted, a regular dispersion of signaling males is the pattern observed in the preferred microhabitats of this species in nature. Offprint requests to: T. Eiriksson     

The timing of calling,movement, and mating in the field crickets Gryllus veletis,G. pennsylvanicus,and G. integer

Summary The temporal relationships between mating, calling, and movement were studied in natural habitats at different densities in the field crickets, Gryllus veletis, G. pennsylvanicus and G. integer. There was variation between and within species, but males generally showed increased calling at dawn followed by a decline during the day. Matings were also more frequent at sunrise in most observations, and male and female movement declined near dawn. Female movement and sexual receptivity are closely related to levels of male calling and movement.     

Risk trading in mating behavior: forgoing anti-predator responses reduces the likelihood of missing terminal mating opportunities

Life history theory predicts that organisms make certain adjustments to their current and future reproductive effort such that fitness is maximized. Moreover, these adjustments may be fine tuned in response to risks of attack by natural enemies. Thus, we may predict that as an organism ages it will accept increasing levels of exposure to predators during mating activities, effectively trading the risk of losing terminal mating opportunities for the risk of predation. We tested this prediction in an acoustic moth, Achroia grisella, in which females orient toward and evaluate males based on their ultrasonic calling song, and both sexes may be vulnerable to predation by insectivorous bats while in flight as well as on the substrate. In the latter situation, singing males and orienting females show silence and arrestment responses, respectively, when presented with synthetic bat echolocation signals broadcast above a threshold amplitude. We found that both males and females become less sensitive to these broadcasts over the course of their brief reproductive periods, 7 and 5 days, respectively. Over the same periods, sensitivity to male song in both males and females remains constant, and relatively little senescence in sexual behavior is observed. These results support the risk trading hypothesis, and they indicate that life history principles may apply over a very short lifespan.     

Growth rate and the cost of calling activity in male carpenter frogs,Rana virgatipes

Summary Energy stress during the breeding season and relationships between calling activity and growth were investigated in male carpenter frogs, Rana virgatipes. This species has a prolonged breeding season of up to three months in Southern New Jersey. Monthly collections made in 1985 revealed that both dry mass and percent body lipid decreased throughout the breeding season but sharply increased at the end of the breeding season. Observations of free-living males showed that small males were more likely to gain mass than large males during the breeding season. All males gained mass at higher rates after the breeding season. A simultaneous record of calling activity and mass change was obtained for 42 males. Males called on 95% of nights, indicating that they rarely ceased their reproductive activities. Small males tended to have low calling efforts and high growth rates. When the effect of initial mass was removed, growth rate was negatively correlated with calling effort among small males. This is the first demonstration of a direct tradeoff between a reproductive activity and growth in an anuran.     

Quantifying Loss of Acoustic Communication Space for Right Whales in and around a U.S. National Marine Sanctuary

The effects of chronic exposure to increasing levels of human‐induced underwater noise on marine animal populations reliant on sound for communication are poorly understood. We sought to further develop methods of quantifying the effects of communication masking associated with human‐induced sound on contact‐calling North Atlantic right whales (Eubalaena glacialis) in an ecologically relevant area (～10,000 km²) and time period (peak feeding time). We used an array of temporary, bottom‐mounted, autonomous acoustic recorders in the Stellwagen Bank National Marine Sanctuary to monitor ambient noise levels, measure levels of sound associated with vessels, and detect and locate calling whales. We related wind speed, as recorded by regional oceanographic buoys, to ambient noise levels. We used vessel‐tracking data from the Automatic Identification System to quantify acoustic signatures of large commercial vessels. On the basis of these integrated sound fields, median signal excess (the difference between the signal‐to‐noise ratio and the assumed recognition differential) for contact‐calling right whales was negative (?1 dB) under current ambient noise levels and was further reduced (?2 dB) by the addition of noise from ships. Compared with potential communication space available under historically lower noise conditions, calling right whales may have lost, on average, 63–67% of their communication space. One or more of the 89 calling whales in the study area was exposed to noise levels ≥120 dB re 1 μPa by ships for 20% of the month, and a maximum of 11 whales were exposed to noise at or above this level during a single 10‐min period. These results highlight the limitations of exposure‐threshold (i.e., dose‐response) metrics for assessing chronic anthropogenic noise effects on communication opportunities. Our methods can be used to integrate chronic and wide‐ranging noise effects in emerging ocean‐planning forums that seek to improve management of cumulative effects of noise on marine species and their habitats. Cuantificación de la Pérdida de Espacio de Comunicación Acústica para Ballenas Francas Dentro y Alrededor de un Santuario Marino Nacional en E. U. A.     

Structure and evolution of song form in the wrens Thryothorus sinaloa and T. felix

Summary The songs of two species of wrens, Thryothorus sinaloa and T. felix, are less similar than previously believed. We have found that song structure, bout patterning, and other aspects of singing behavior are widely divergent between the two species, which are sympatric over much of their ranges. The nature of these differences supports the hypothesis of contrast reinforcement in bird song. Further evidence is provided by essentially parallel differences in another pair of sympatric species, T. pleurostictus and T. maculipectus. A cluster analysis of song form of the above four species plus one other, T. ludovicianus, suggests that those species which are allopatric are the most similar in song form, while sympatric species are the most dissimilar. We relate various features of singing behavior in the two species to some aspects to their ecology, and examine relationships among various song parameters within the genus Thryothorus. The parallel differences in pairs of sympatric Thryothorus in structure of song units, temporal organization of song, and duetting, along with the close correlation of groups of song parameters in the five species of Thryothorus leads us to propose that strategies or adaptive complexes of singing characteristics may exist in the genus.