Early singers attend to conspecific but not heterospecific behavioural cues at dawn

During the dawn chorus songbirds initiate singing activities just prior to sunrise. Environmental factors affect the timing of the dawn chorus, but relatively little is known about how behavioural cues influence chorus timing. We assessed whether early playback of conspecific and heterospecific song influenced chorus start times in two early‐singing temperate species, hermit thrush Catharus guttatus and veery C. fuscescens. We used arrays of GPS time‐synchronized recorders to record the dawn chorus at 30 locations. In each location, thirty minutes prior to natural chorus initiation, we broadcast: 1) an early singing species (hermit thrush or veery), 2) a late singing species (black‐throated green warbler Setophaga virens or Nashville warbler Oreothlypis ruficapilla), 3) a noise control, and 4) also recorded without playback (silent control). Playback treatments were separated by two‐three days at each location. Both hermit thrush and veery sang significantly earlier in response to conspecific playback compared to noise or silent controls. Both species also sang earlier in response to conspecific playback compared to the day before or the day after playback. Neither species sang earlier in response to heterospecific playback (regardless of whether the broadcast species was a naturally early or late song initiator) compared to noise or silent controls. Our results support the theory that the dawn chorus is a communication network where individuals attend primarily to conspecific cues; heterospecific song appears to have minimal impacts on the chorus start times of the two early‐singing species we investigated.     

<Emphasis Type="Italic">Drosophila</Emphasis> pupation behavior in the wild

We investigated pupa distributions of D. simulans, D. buzzatii, D. melanogaster, D. immigrans and D. hydei on a number of natural breeding sites. Pupae of all five species showed aggregated distributions, which prompted us to examine these aggregations in a more detail for two species that commonly co-occur in breeding sites, D. simulans and D. buzzatii. We found that pupae of both species tend to be aggregated in conspecific clusters. Subsequent experiments revealed that both species are attracted to the odors of other larvae, though only D. buzzatii differentiated between conspecifics and heterospecifics (they preferred conspecific). Furthermore, third instar larvae of both species preferred more alkaline substrates. Altogether, our results demonstrate that Drosophila species form conspecific pupa aggregations in natural breeding sites, and that pupation site selection depends on interactions among conspecific and heterospecific larvae and on chemical characteristics of the breeding sites.     

Differential Response to Interspecific and Intraspecific Signals Amongst Chickadees

Black‐capped chickadees (Poecile atricapillus) and mountain chickadees (P. gambeli) have a similar vocal repertoire and share many other life history traits; yet, black‐capped chickadees are socially dominant to mountain chickadees where populations overlap. Previous research suggested that in contact zones, both species respond weakly to heterospecific songs during the breeding season, and have suggested minimal interspecific competition. However, both black‐capped and mountain chickadees discriminate between conspecific and heterospecific chick‐a‐dee calls, suggesting attention is paid to interspecific signals. We compared the responses of both black‐capped and mountain chickadees to conspecific and heterospecific chick‐a‐dee calls during the winter, when both species compete for the same food resources. We conducted an aviary playback experiment exposing both species to playback composed of heterospecific and conspecific chick‐a‐dee calls, which had been recorded in the context of finding food sources. Responses from the tested birds were measured by recording vocalizations and behaviour. Black‐capped chickadees responded significantly more to conspecific than to heterospecific stimuli, whereas the subordinate mountain chickadees responded to both mountain and black‐capped chickadee calls. Based upon the reactions to playbacks, our results suggest these two closely related species may differ in their perception of the relative threat associated with intra‐ versus interspecific competitors.     

Phonotaxis to male’s calls embedded within a chorus by female gray treefrogs, Hyla versicolor

During the reproductive season, male Hyla versicolor produce advertisement calls to attract females. Females exhibit phonotaxis and approach the individual callers, resulting in amplexus. For frogs that call from dense choruses, the extent to which and the range from which a male’s advertisement call within a chorus can be heard by a receptive female leading to phonotaxis is unclear. We investigated females’ responses to natural choruses in the field and found that they were attracted and showed directed orientation to breeding choruses at distances up to 100 m. To assess the role of acoustic cues in the directed orientation, we conducted acoustic playback experiments in the laboratory using conspecific call and noise as stimuli, as well as chorus sounds (that contained calls from a focal male) recorded at various distances, all played at naturalistic intensities. Using two response metrics (females’ normalized response times and their phonotaxis trajectories) we found that, unlike the field experiments, females oriented and were attracted to chorus sounds from 1 to 32 m only, but not from >32 m, or to band-limited noise. Possible reasons for the observed difference in phonotaxis behavior in the two experimental conditions were discussed.     

Sexual differentiation and seasonal variation in response to conspecific and heterospecific acoustic signals

Interspecific territoriality is frequently reported between closely related species; however, few studies have demonstrated interspecific territoriality between distantly related species living in sympatry. We conducted playback experiments to investigate territorial behaviour in male and female White‐bellied Wrens (Uropsila leucogastra) in response to simulated conspecific and heterospecific intruders during the breeding and non‐breeding seasons. We explored whether heterospecific songs of the Happy Wren (Pheugopedius felix), a distantly related species and ecological competitor, elicited antagonistic responses from focal White‐bellied Wrens, and whether such responses differed between the sexes. We also examined whether male and female responses to conspecific and heterospecific rivals varied with season. We found that male White‐bellied Wrens always responded to conspecific song, and responded significantly more to heterospecific song compared to a control stimulus (Tropical Parula, Setophaga pitiayumi). In contrast, although female White‐bellied Wrens responded strongly to conspecific song, their response to heterospecific song did not differ significantly from the control stimulus. The proportion of males that responded to heterospecific songs and the proportion of females that responded to conspecific songs varied seasonally, showing significantly lower responses during the breeding season. The intense responses of male White‐bellied Wrens to playback of heterospecific songs suggest that they recognise ecological competitors based on their vocal signals. Furthermore, the decrease in agonistic interactions during the breeding season is in line with the hypothesis that aggressive behaviour may be detrimental to reproductive and parental activity, and the hypothesis that heterospecific animals pose less of a threat during the breeding season.     

Who is in the neighborhood? Conspecific and heterospecific responses to perceived density for breeding habitat selection

Theoretical models of habitat selection often incorporate negative density dependence. Despite strong negative density‐dependent effects on habitat selection, more recent studies indicate that animals settle near members of their own (conspecific) and other species (heterospecific) when selecting habitat with social cues. Social cue use for habitat selection is particularly common among songbirds, but few studies have investigated if songbirds use social cues to assess conspecific or heterospecific density (as opposed to just presence/absence) when making settlement decisions. We conducted a playback experiment to evaluate if yellow warblers (Setophaga petechia) and willow flycatchers (Empidonax traillii), two potential competitors for breeding habitat, use social cues to assess density (conspecific for warblers and heterospecific for flycatchers) when selecting breeding locations at two spatial scales. We simulated yellow warbler density to be high or low at multiple treatment plots (3.14 ha) with song playback and then evaluated settlement decisions by comparing yellow warbler and willow flycatcher abundances across plots (broad‐scale habitat selection) and individual space use within plots (fine‐scale territory establishment). Yellow warbler density treatments did not affect habitat selection by yellow warblers at the broad scale, but caused individuals to cluster territories at high‐density treatments. Willow flycatchers were most abundant at high‐density treatment plots, but yellow warbler density treatments did not affect territory locations. The results indicate that perceived density affects the habitat selection process for both conspecifics and heterospecifics.     

MATE CHOICE OCCURS ONLY IN SMALL CHORUSES OF PAINTED REED FROGS HYPEROLIUS MARMORATUS

ABSTRACT

In two-choice discrimination experiments, females of Hyperolius marmoratus preferred the calls of lower frequency of the pair of stimuli. This preference was not shown in mating patterns observed in natural choruses, but is when females are phonotactically orienting in small choruses in an experimental enclosure. With an increase in chorus size, the mating pattern shifts from size-based, non-random (with some evidence of size-assortative) mating to random mating. This is the first time that frequency-based mate-choice by female anurans has been associated with chorus size, and hence with the sonic complexity of the acoustic environment.     

Interspecific Response to Playback of Bird Song

Responses to bird song have usually only been studied at the intraspecific level. I experimentally tested whether playback of the song of the black wheatear Oenanthe leucura in an area in S Spain resulted in responses from conspecifics as well as heterospecific birds by comparing the numbers of individuals singing before and after playback. The number of singing male black wheatears increased considerably, but also the number of singing males of five other passerine species increased significantly. The heterospecific response to playback may be due (1) to interspecific territoriality, (2) to black wheatear song signalling the absence of predators, or (3) to heterospecifics confusing the species-identity of the singer. The second alternative is considered more likely, since an ecologically wide array of species increased their song rate following playback. The conspicuous dawn (and dusk) chorus of bird song may be augmented by social facilitation due to the singing of conspecifics as well as heterospecifics.     

Spatial Nest‐Settlement Decisions in Digger Wasps: Conspecifics Matter more than Heterospecifics and Previous Experience

Research into the driving forces behind spatial arrangement of wasp nests has considered abiotic environmental factors, but seldom investigated attraction or repulsion towards conspecifics or heterospecifics. Solitary female digger wasps (Hymenoptera) often nest in dense aggregations, making these insects good models to study this topic. Here, we analysed the nesting patterns in an area shared by three species of the genus Bembix, in a novel study to discover whether female wasps are attracted to or repulsed by conspecific nests, heterospecific nests or their own previously established nests when choosing nest‐digging locations. Early in the season, each species showed a clumping pattern of nests, but later in the season, a random distribution of nests was more common, suggesting an early conspecific attraction. Such behaviour was confirmed by the fact that females started building their nests more frequently where other females of their species were simultaneously digging. The distances between subsequent nests dug by individual females were shorter than those obtained by random simulations. However, this pattern seemed to depend on the tendency to dig close to conspecifics rather than remain in the vicinity of previous nests, suggesting that females' experience matters to future decisions only on a large scale. Nesting patches within nest aggregations largely overlapped between species, but the nests of each species were generally not closer to heterospecific nests than expected by chance, suggesting that females are neither repulsed by, nor attracted to, congenerics within nest aggregations. A role of the spatial distribution of natural enemies on the observed nesting patterns seemed unlikely. Bembix digger wasp nest aggregations seem thus to be primarily the result of female–female attraction during nest‐settlement decisions, in accordance with the ‘copying’ mechanisms suggested for nesting vertebrates.     

Frog breeding pond selection in response to predators and conspecific cues

Predators are a major influence on the breeding site selection decisions of anurans. Many species actively avoid breeding in habitat with predators when given the choice between predator and predator‐free sites. However, certain factors such as site fidelity or conflicting cues may preclude avoidance behavior. We conducted two experiments examining how western chorus frogs, Pseudacris triseriata, respond to predators, western mosquitofish, Gambusia affinis, using an array of artificial ponds located at two field sites. In one experiment, we added G. affinis to half of our experimental ponds and monitored subsequent colonization by frogs. We found that frogs laid significantly fewer eggs in ponds with fish compared to fishless ponds. In another experiment, we introduced an additional cue to complicate the decision‐making process and monitored colonization of ponds in response to treatments of conspecific breeding cues only (eggs), predators (G. affinis) only, and conspecific cues and predators. We found no significant differences in number of eggs deposited among these three treatments. Based on these results, P. triseriata does not always exhibit complete avoidance of fish predators, and avoidance may vary based on factors such as site fidelity or dispersal costs. This study represents a step toward understanding how multiple biotic factors at a breeding pond may influence anuran site selection behavior in the field.     

Modelling the effects of chorus species composition and caller density on acoustic masking interference in multispecies choruses of crickets and katydids

Natural multispecies acoustic choruses such as the dusk chorus of a tropical rain forest consist of simultaneously signalling individuals of different species whose calls travel through a common shared medium before reaching their ‘intended’ receivers. This causes masking interference between signals and impedes signal detection, recognition and localization. The levels of acoustic overlap depend on a number of factors, including call structure, intensity, habitat-dependent signal attenuation and receiver tuning. In addition, acoustic overlaps should also depend on caller density and the species composition of choruses, including relative and absolute abundance of the different calling species. In this study, we used simulations to examine the effects of chorus species relative abundance and caller density on the levels of effective heterospecific acoustic overlap in multispecies choruses composed of the calls of five species of crickets and katydids that share the understorey of a rain forest in southern India. We found that on average species-even choruses resulted in higher levels of effective heterospecific acoustic overlap than choruses with strong dominance structures. This effect was found consistently across dominance levels ranging from 0.4 to 0.8 for larger choruses of forty individuals. For smaller choruses of twenty individuals, the effect was seen consistently for dominance levels of 0.6 and 0.8 but not 0.4. Effective acoustic overlap (EAO) increased with caller density but the manner and extent of increase depended both on the species' call structure and the acoustic context provided by the composition scenario. The Phaloria sp. experienced very low levels of EAO and was highly buffered to changes in acoustic context whereas other species experienced high EAO across contexts or were poorly buffered. These differences were not simply predictable from call structures. These simulation-based findings may have important implications for acoustic biodiversity monitoring and for the study of acoustic masking interference in natural environments.     

Long‐Distance Calling by the Willow Tit,Poecile montanus,Facilitates Formation of Mixed‐Species Foraging Flocks

The occurrence of mixed‐species foraging flocks is a worldwide phenomenon in terrestrial bird communities. Previous studies suggest that individuals participating in flocks might derive benefits in terms of improved feeding efficiency and/or reduced risk of predation. However, very little is known about how individuals establish mixed‐species flocks. Here, I provide the first experimental evidence that long‐distance calling by the willow tit, Poecile montanus, facilitates the establishment of mixed‐species flocks at a foraging patch. Observations at experimental foraging patches showed that willow tits that find a food source produce long‐distance calls, particularly when they are isolated from conspecific flockmates. The probability of long‐distance calling was negatively correlated with the number of heterospecific foraging individuals near the food source. A playback experiment confirmed that calls attract both conspecific and heterospecific members of foraging flocks. This study demonstrates that willow tits use long‐distance calls to attract conspecific flockmates to foraging patches, and these calls can also facilitate the formation of mixed‐species flocks on patches.     

Ravens adjust their antipredatory responses to con‐ and hetero‐specific alarms to the perceived threat

Heterospecific alarm calls are typically found in situations where multiple species have a common predator. In birds, they are particularly common in mixed mixed‐species flocks. In species with highly developed social and cognitive abilities like corvids, there is the potential for differential responses to heterospecific vs. conspecific calls according to the riskiness of the habitat. We tested the responses of free‐ranging ravens (Corvus corax) to conspecific alarm calls and compared them to heterospecific alarm calls of jackdaws (Corvus monedula). We observed the proportion of ravens leaving the feeding site after the con‐ or hetero‐specific playback was presented in a situation of low threat (wild boar—Sus scrofa enclosure) and high threat of predation (wolf—Canis lupus enclosure). We show that ravens responded to conspecific calls more intensively at the wolves than at the wild boar, but the response to conspecific calls was in both enclosures stronger than to the control (great tit—Parus major song). The response to the heterospecific alarm was also stronger in the wolves’ enclosure, but it did not differ from control in the wild boar enclosure. These findings suggest that ravens are aware of the meaning of the jackdaw alarm calls, but they respond to it only in a situation of high predatory threat (wolves are present). In the wild boar enclosure, the ravens probably consider jackdaws warning against some other predator, very probably harmless to ravens. This interpretation requires further testing, as both enclosures differ also in respect to other parameters like food quality and shelter availability.     

Acoustic Preference of Frog‐Biting Midges (Corethrella spp) Attacking Túngara Frogs in their Natural Habitat

In many animals, males aggregate to produce mating signals that attract conspecific females. These leks, however, also attract eavesdropping predators and parasites lured by the mating signal. This study investigates the acoustic preferences of eavesdroppers attracted to natural choruses in a Neotropical frog, the túngara frog (Engystomops pustulosus). In particular, we examined the responses of frog‐biting midges to natural variation in call properties and signaling rates of males in the chorus. These midges use the mating calls of the frogs to localize them and obtain a blood meal. Although it is known that the midges prefer complex over simple túngara frog calls, it is unclear how these eavesdroppers respond to natural call variation when confronted with multiple males in a chorus. We investigated the acoustic preference of the midges using calling frogs in their natural environment and thus accounted for natural variation in their call properties. We performed field recordings using a sound imaging system to quantify the temporal call properties of males in small choruses. During these recordings, we also collected frog‐biting midges attacking calling males. Our results revealed that, in a given chorus, male frogs calling at higher rates and with higher call complexity attracted a larger number of frog‐biting midges. Call rate was particularly important at increasing the number of midges attracted when males produced calls of lower complexity. Similarly, call complexity increased attractiveness to the midges especially when males produced calls at a low repetition rate. Given that female túngara frogs prefer calls produced at higher repetition rates and higher complexity, this study highlights the challenge faced by signalers when increasing attractiveness of the signal to their intended receivers.     

Vocal Strategies in Confronting Interfering Sounds by a Frog from the Southern Temperate Forest,Batrachyla antartandica

Animals are communicating by sound face interference from biotic and abiotic sources. Contrasting strategies have been reported in different taxa in the presence of prolonged noises, but in particular, interactions among acoustically active species have been studied to a very limited extent. In addition, reactions of a single species to interferences having contrasting structural patterns have not been explored systematically. The vocal responses of 16 male frogs Batrachyla antartandica from the temperate austral forest in Chile were tested with conspecific calls and with the calls of two sympatric species: B. taeniata and B. leptopus, broadcast at amplitudes of 73, 79, 85, 91, and 97 dB peak sound pressure level (SPL). Also, the vocal activity of the subjects during exposure to a 3‐min continuous broadband noise presented at 67 dB root mean square (RMS) SPL was monitored. The subjects gave significantly higher responses to the conspecific relative to the heterospecific calls but increased their vocal output in the presence of continuous noise. The preference of vocal responses for the conspecific relative to heterospecific signals bears a general resemblance to those exhibited by B. taeniata in a previous study and potentially contributes to the typical segregation in conspecific choruses observed in areas where these frogs breed in sympatry. Such lack of vocal responses to heterospecific stimuli contrasts with the increase in vocal activity during exposure to continuous noise lacking a temporal fine structure and points to identify different strategies in confronting interferences of diverse origin.     

Reproductive isolation inProkelisia planthoppers (Homoptera: Delphacidae): Acoustic differentiation and hybridization failure

Males and females of Prokelisia marginata (Van Duzee) and Prokelisia dolusWilson communicate through substrate-transmitted vibrations. The acoustic signals (attraction and courtship calls) of these planthoppers are effective in mate location, attraction, and mate choice. Attraction calls are structurally distinct for both species and differ in pulse type, pulse repetition rate, and pulse duration. Using playback of prerecorded calls, individuals discriminated between conspecific and heterospecific signals. Depending on the sex and species, response calls were produced three to eight times more frequently to conspecifics than to heterospecifics. However, acoustic signals alone did not explain reproductive isolation and hybridization failure in these two congeners. Some heterospecific pairs called, courted, and attempted to join genitalia, but no connections were successful and no progeny were produced. Thus, acoustic behavior is not a guaranteed premating isolating mechanism in no-choice situations. Other courtship behaviors and possibly morphological differences in genitalia also contributed to their isolation. Females displayed a variety of rejection behaviors to conspecific and heterospecific males, suggesting that sexual selection (female choice), in addition to species recognition, may be an important force in the evolution of the acoustic signals of planthoppers. Although signal structure was not dependent on wing form (planthoppers exhibit wing dimorphism), the age when females first began to call was related to wing form. Brachypterous (flightless) females of both species began calling early in adult life (day 2), whereas macropterous (migratory) females began calling later in adult life (day 6). This pattern is consistent with the oogenesis-flight syndrome, in which reproductive maturity is delayed until after migration occurs.     

The Sounds of Silence as an Alarm Cue in Túngara Frogs,Physalaemus pustulosus

In many species males vocally advertise for mates in choruses and these choruses serve as acoustic beacons to conspecific females as well as to eavesdropping predators and parasites. Chorusing will often cease in response to disturbances, such as the presence of predators. In some cases the cessation is so rapid and over such a large area that it seems improbable that males are all responding directly to the same local disturbance. Here, we demonstrate experimentally in Neotropical túngara frogs, Physalaemus pustulosus, that the cessation of calling by males spreads rapidly through the chorus. The cessation of chorusing in response to the cessation of playbacks of three calling males is more effective in inducing chorus cessation than is the cessation of one male calling. When three males are calling, the cessation of complex calls is more effective in inducing chorus cessation than simple calls. There is no main effect on whether the final call of the male is complete or is interrupted. We thus conclude that the sudden lack of signals—the ‘sounds of silence’—becomes an alarm cue that can explain the rapid cessation of choruses that are common in many chorusing species.     

Wild fledgling tits do not mob in response to conspecific or heterospecific mobbing calls

Mobbing, where birds harass a predator through a combination of vocalizations and stereotyped behaviours, is an effective anti-predator behaviour for many species. Mobbing may be particularly important for juveniles, as these individuals are often more vulnerable than adults. Although the component behaviours of mobbing are often considered to be un-learned, there are few confirmatory data, and the developmental trajectory of mobbing is unknown. In this study, we tested whether conspecific or heterospecific mobbing calls initiated mobbing behaviour in juvenile Blue Tits Cyanistes caeruleus. We located wild adult and recently fledged juvenile Blue Tits and presented them with playback recordings of adult conspecific (Blue Tit) and heterospecific (Great Tit Parus major) mobbing alarm calls. Although adult birds readily mob in response to these types of playbacks, juveniles did not exhibit characteristic mobbing behaviour. Some juveniles did, however, exhibit individual components of mobbing behaviour found in mobbing, despite not producing adult-like mobbing behaviour in response to either conspecific or heterospecific playback. These results suggest that, although birds might be capable of mobbing as juveniles, the associations between the non-vocal stereotyped mobbing behaviours and mobbing calls may be learned.     

Dynamics of periodical cicada chorus centers (Homoptera: Cicadidae:Magicicada)

Mating aggregations of three species of periodical cicadas were monitored during the emergence of Brood XIX at a 16-ha study site in northwest Arkansas, May–June 1985. Magicicada tredecassiniappeared first and formed the most choruses. M. tredecimand M. tredeculachoruses formed next, and M. tredeculachoruses outnumbered those of M. tredecim.Of the 268 choruses seen, 84% were composed of M. tredecassini. M. tredecassiniwere often found chorusing in the same trees with the other two species. Such multispecies mating aggregations apparently are unique to periodical cicadas. Choruses were dynamic with respect to their locations and durations. Initially, choruses were located near areas of high cicada emergence densities. One week later, cicadas chorused in trees throughout the forest and at the forest edge. Many choruses were seen only once at a location. Although cicadas chorused for almost 4 weeks, individual choruses persisted only approximately 8 days, on average. Sound intensities under chorus centers ranged from 50 to 80 dh and were correlated with arena sizes during times of peak chorus activity. No distinct habitat preferences of the three species were observed, however, the tree species used by chorusing cicadas differed among the species.     

Neighbor Call Amplitude Influences Aggressive Behavior and Intermale Spacing in Choruses of the Pacific Treefrog (Hyla regilla)

Male Pacific treefrogs (Hyla regilla) aggregate in choruses during the breeding season. Within these choruses frogs distribute themselves nonrandomly. This study tested the hypothesis that the amplitude of neighbors' calls serves as a proximate cue in regulating the spacing of males in choruses, and that this is mediated by the incidence of aggressive-encounter calls by resident males. The amplitude of neighbors' calls showed little interindividual variation. Advertisement calls were played to males at three amplitudes that spanned the range of neighbor-call amplitudes measured between pairs of frogs. At playback amplitudes corresponding to the minimum neighbor-call amplitudes observed in choruses, frogs gave predominantly advertisement calls and few aggressive-encounter calls in response. As the playback amplitude was increased, subjects progressively decreased the number of advertisement calls and increased the number of encounter calls that they produced in response. The total number of calls (advertisement + encounter) given in response did not vary with playback amplitude. Intruders were likely to move away when a resident male gave encounter calls. In this way neighbor-call amplitude regulates intermale spacing in choruses.