Soundscape diversity in the Great Barrier Reef: Lizard Island,a case study

Passive acoustic monitoring can provide valuable information on coral reefs, and examining the acoustic attributes of these ecosystems has the potential to provide an insight into their status and condition. From 2014 to 2016, a series of underwater recordings were taken at field sites around Lizard Island in the Great Barrier Reef, Australia. Six individual fish choruses were identified where each chorus displayed distinct acoustic characteristics. Choruses exhibited diurnal activity and some field sites displayed consistently higher diversity of choruses and levels than others, suggesting that particular locations are important aggregation areas for soniferous fish species. During peak activity, choruses were a prominent component of reef soundscapes, where received levels of a chorus reached upwards of 120 dB re 1μPa rms over the 450–650 Hz band, equating to a 40 dB increase above ambient noise levels of ≈80 dB re 1μPa rms. Three out of the six detected choruses exhibited spectral and temporal characteristics similar to choruses previously documented at these sites and elsewhere, produced by planktivorous fish species. Three of these choruses appear to be undocumented and could hold information on the presence, abundance and dispersal patterns of important fish species, which may have potential long-term management applications. Future research should focus on extricating the temporal patterns associated with bioacoustic activity and determining the potential environmental drivers of biological choruses. Additionally, developing appropriate techniques for direct identification of vocalizing species would strongly increase the management applicability of passive acoustic monitoring.     

Fish choruses off Port Hedland,Western Australia

Australian waters are home to a number of vocal species of fish. Cataloguing the acoustic characteristics and temporal patterns of choruses and their locations can provide significant information for long-term monitoring of vocal fishes and their ecosystems. In coastal waters off Port Hedland, Western Australia, two seafloor positioned sea-noise loggers, located 21.5 km apart in 8 and 18 m of water, recorded for an 18-month period. Numerous sound sources were detected, including mooring and vessel noise, humpback whale song and a large variety of fish signal types. Seven fish choruses were identified, occurring predominantly between late spring and early autumn (wet season) and displaying energy from 50 Hz to >4 kHz. Many of these choruses exhibited acoustic characteristics similar to choruses previously reported elsewhere, for some of which the source species or families have been identified. Distinct diurnal patterns in the choruses were observed, associated with sunrise or sunset and in some cases, both. While choruses were predominantly recorded on different days, there were at total of 80 days when more than one chorus was present at the same site. Some pairs of choruses present on the same day exhibited various combinations of temporal and frequency partitioning, while others displayed predominant overlap in both spaces.     

Inter-male Spacing and Aggression in African Painted Reed Frogs,Hyperolius marmoratus

Inter-male spacing and the role of aggression in the maintenance of spatial organization was examined in choruses of Hyperolius marmoratus males housed in a semi-natural enclosure. The effects of chorus size on spacing patterns, nearest neighbour distances and male aggressive behaviour were examined. Males were found to reduce nearest neighbour distances as chorus size increased. This was accompanied by a shift in spacing pattern from even in small choruses (5–9 males) to random in choruses of 10 to 13 males. In choruses of 14, spacing patterns were once again even. The reason for these shifts is unclear but may reflect a sudden, rather than gradual increased tolerance by individuals to high neighbour call intensity as space to call from becomes more limited. The level of male aggression was influenced by chorus size and the time of night. In general, more aggressive interactions occurred in high density choruses. However, this did not translate into higher levels of individual male aggression as density increased. Individual male aggression was high in early evening choruses and declined to a minimum at peak chorusing time. High levels of aggression during early evening may reflect the establishment of calling sites by males, while the drop in aggression at peak chorusing time may occur in response to the presence of females in the chorus at this time or as a consequence of masking of neighbours calls.     

Fine‐scale habitat use in Indo‐Pacific humpback dolphins,Sousa chinensis,may be more influenced by fish rather than vessels in the Pearl River Estuary,China

Shifts in habitat use and distribution patterns in dolphins are often concerns that can result from habitat degradation. We investigated how potential changes to a habitat from human activity may alter dolphin distributions within Lingding Bay in the Pearl River Estuary, China, by studying the relationship between fish choruses, vessel presence and Indo‐Pacific humpback dolphin (Sousa chinensis) detection rates. Analyses revealed temporal and spatial variation within fish choruses, vessel presence and dolphin detection rates. After accounting for any temporal autocorrelation, correlations between fish choruses and dolphin detection rates were also found; however, no relationship between fish choruses and vessel presence or dolphin detection rates and vessel presence were observed. Furthermore, fewer dolphins were detected at sites where fish activity was less intense. Thus fish activity, rather than vessels, may be a key factor influencing the distribution of the dolphins within the estuary. These findings emphasize the risk of potential shifts in habitat use for Indo‐Pacific humpback dolphins due to detrimental changes to prey availability and dolphin feeding grounds from human activity, such as overfishing and coastal developments, within the estuary. This is a critical conservation issue for this dolphin population that is facing intense anthropogenic pressure.     

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.     

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.     

A rain forest dusk chorus: cacophony or sounds of silence?

A rain forest dusk chorus consists of a large number of individuals of acoustically communicating species signaling at the same time. How different species achieve effective intra-specific communication in this complex and noisy acoustic environment is not well understood. In this study we examined acoustic masking interference in an assemblage of rain forest crickets and katydids. We used signal structures and spacing of signalers to estimate temporal, spectral and active space overlap between species. We then examined these overlaps for evidence of strategies of masking avoidance in the assemblage: we asked whether species whose signals have high temporal or spectral overlap avoid calling together. Whereas we found evidence that species with high temporal overlap may avoid calling together, there was no relation between spectral overlap and calling activity. There was also no correlation between the spectral and temporal overlaps of the signals of different species. In addition, we found little evidence that species calling in the understorey actively use spacing to minimize acoustic overlap. Increasing call intensity and tuning receivers however emerged as powerful strategies to minimize acoustic overlap. Effective acoustic overlaps were on average close to zero for most individuals in natural, multispecies choruses, even in the absence of behavioral avoidance mechanisms such as inhibition of calling or active spacing. Thus, call temporal structure, intensity and frequency together provide sufficient parameter space for several species to call together yet communicate effectively with little interference in the apparent cacophony of a rain forest dusk chorus.     

CHORUS HOWLING BY WOLVES: ACOUSTIC STRUCTURE,PACK SIZE AND THE BEAU GESTE EFFECT

ABSTRACT

A variety of structural parameters were measured from wolf choruses recorded in the Superior National Forest, Minnesota, USA. Mean duration of 60s did not vary with pack size or composition. Packs replied to simulated howling after an average of 40s, often interrupting the stimulus howls. Choruses began with simply-structured howls, which became increasingly modulated as the chorus progressed. Little difference in mean fundamental frequency or other howl parameters was found among the choruses from packs of various sizes and compositions. In particular, choruses produced by single adult pairs did not differ from those of larger packs accompanied by pups. The lack of relationship between chorus parameters and pack size or composition indicates there is little useful information concerning a pack's size to be found in its chorus howling.

The observation that chorus howling by adult pairs is often perceived as that of larger groups with pups suggests that chorus structure has evolved to exaggerate the apparent size of the pack, especially those newly-established or otherwise reduced in number. If so, wolf howling choruses may represent a mammalian example of the Beau Geste effect, made particularly viable because of the relative immunity of the signal to probing.     

Female preference for temporal features of vocalization in the Japanese treefrog,Rhacophorus arboreus

Female preference for the different temporal features of advertisement calls by males was studied in the foam-nesting treefrog,Rhacophorus arboreus. Field observations showed that females were attracted to males making more calls, calling in more bouts of vocalization and leading choruses. In phonotactic experiments in the laboratory, however, females were attracted to vocalizations with more total notes while they were not attracted to chorus leaders. alphabetical order     

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.     

Owl dusk chorus is related to the quality of individuals and nest‐sites

Dawn and dusk choruses represent one of the most investigated topics in avian vocal behaviour, but their underlying basis remains unclear. As with the dawn chorus in passerines, dusk chorus in owls seems to support the mate and rival assessment hypothesis and happens during the most constraining period, as individuals have not yet fed and, under the handicap principle, dusk chorus is likely to reveal inter‐individual differences in competitive ability, body condition and/or habitat quality. Here, a study of vocal displays at dusk of 14 Eurasian Eagle Owls Bubo bubo revealed a temporal succession in the order in which males began their vocalizations. The vocalization order appeared to be related to both the quality of the nesting territory (based upon mean number of fledged young and proportion of rats in the diet) and the male's individual quality, as revealed by haematocrit values and the brightness of the white throat patch.     

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.     

Self‐organizing cicada choruses respond to the local sound and light environment

1. Periodical cicadas exhibit an extraordinary capacity for self‐organizing spatially synchronous breeding behavior. The regular emergence of periodical cicada broods across the United States is a phenomenon of longstanding public and scientific interest, as the cicadas of each brood emerge in huge numbers and briefly dominate their ecosystem. During the emergence, the 17‐year periodical cicada species Magicicada cassini is found to form synchronized choruses, and we investigated their chorusing behavior from the standpoint of spatial synchrony.
2. Cicada choruses were observed to form in trees, calling regularly every five seconds. In order to determine the limits of this self‐organizing behavior, we set out to quantify the spatial synchronization between cicada call choruses in different trees, and how and why this varies in space and time.
3. We performed 20 simultaneous recordings in Clinton State Park, Kansas, in June 2015 (Brood IV), with a team of citizen‐science volunteers using consumer equipment (smartphones). We use a wavelet approach to show in detail how spatially synchronous, self‐organized chorusing varies across the forest.
4. We show how conditions that increase the strength of audio interactions between cicadas also increase the spatial synchrony of their chorusing. Higher forest canopy light levels increase cicada activity, corresponding to faster and higher‐amplitude chorus cycling and to greater synchrony of cycles across space. We implemented a relaxation‐oscillator‐ensemble model of interacting cicadas, finding that a tendency to call more often, driven by light levels, results in all these effects.
5. Results demonstrate how the capacity to self‐organize in ecology depends sensitively on environmental conditions. Spatially correlated modulation of cycling rate by an external driver can also promote self‐organization of phase synchrony.

     

Simulated chorus attracts conspecific and heterospecific Amazonian explosive breeding frogs

In tropical regions, some anuran species breed "explosively", reproducing in massive and highly diverse aggregations during a brief window of time. These aggregations can serve as acoustic beacons, attracting other anurans toward seasonal ponds. We hypothesize that conspecific and heterospecific calls play a role in navigation toward ponds and synchronization of reproduction among species. We simulated a chorus of two species (Trachycephalus coriaceus and Chiasmocleis shudikarensis) with contrasting call characteristics (low‐frequency vs. high‐frequency) and reproductive strategies (strict pond breeder vs. opportunistic) near known explosive breeding sites. We predicted that choruses of T. coriaceus are more attractive to heterospecifics than of C. shudikarensis because the first provides a more reliable indicator of a suitable breeding pond and a better long‐distance signal. We found that both choruses attracted conspecific frogs to the playback outside a natural breeding event. As predicted, heterospecifics were attracted only by low‐frequency calls of T. coriaceus that breeds exclusively in large ponds, but not by higher frequency calls of C. shudikarensis that also breeds in small pools not suitable for other species. Our study presents the first experimental evidence that tropical explosive breeding anurans are attracted to conspecific and heterospecific choruses. The contrasting effect of the playback of the two species on heterospecifics suggests that the attractive effect of a chorus depends on the reproductive strategy of both the sender and the receiver. Given the abundance and diversity of communities in tropical ecosystems, the use of heterospecific acoustic cues may prove widespread and requires further investigation. Abstract in Portuguese is available with online material.     

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.     

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.     

Female Iberian midwife toads (Alytes cisternasii) use call order to reach particular males in dense choruses

Call-timing mechanisms among anuran males calling within choruses are frequent, and previous studies have indicated that midwife toad females respond preferentially to these mechanisms. The results of multi-speaker tests performed here with midwife toads suggest that the temporal order of male calls within a dense chorus can determine the ability of females to locate the most attractive calls (in this instance the low-frequency calls emitted by larger males). When call emission was regular in the multi-speaker tests, most females chose the most attractive option (i.e., the speaker emitting the lower-frequency call). When call emission was not regular, however, most females failed to reach the most attractive option, selecting the speaker that emitted immediately after that emitting the most attractive call. These results support the idea that there may be benefits for smaller males that emit their less attractive calls close to those of larger, more attractive males in dense choruses. Less attractive males could exploit the attractiveness of nearby larger males calls by alternating their calls in an unordered sequence to reach receptive females.     

Mate sampling by female barking treefrogs (Hyla gratiosa)

Despite intense interest in mate choice, relatively little isknown about how individuals sample prospective mates. Indeed,a key issue is whether females sample males or simply matewith the first male encountered. We investigated mate samplingby female barking treefrogs (Hyla gratiosa). Females choosingmates in natural choruses did not move between males but insteadmated with the first male they approached closely. Most femalesmated with the male closest to them at the start of their mate-choiceprocess, and females were more likely to mate with the closest male when the distance to other males was large. These observationsare consistent with the hypothesis that females do not samplepotential mates but instead mate with the first male they distinguishfrom the rest of the chorus. To test this initial detectionhypothesis, we conducted a playback experiment in which weoffered females a choice between two calls, one of which was detectable above the background chorus sound at the female'srelease point, and one of which became detectable only as femalesmoved toward the initially detectable call. Females did notprefer the initially detectable call, thus ruling out the initialdetection hypothesis and implicating sampling of potentialmates by females. Based on the behavior of females in natural choruses, we hypothesize that females approach the chorus, moveto locations where they are able to detect the calls of severalmales simultaneously, and choose a mate from among these malesat some distance from the males. Such simultaneous samplingmay be common in lekking and chorusing species, which havebeen the subjects of many studies of sexual selection.