A framework for the quantification of soundscape diversity using Hill numbers

1. Soundscape studies are increasingly common to capture landscape-scale ecological patterns. Yet, several aspects of soundscape diversity quantification remain unexplored. Although some processes influencing acoustic niche usage may operate in the 24h domain, most acoustic indices only capture the diversity of sounds co-occurring in sound files at a specific time of day. Moreover, many indices do not consider the relationship between the spectral and temporal traits of sounds simultaneously. To provide novel insights into landscape-scale patterns of acoustic niche usage at broader temporal scales, we present a workflow to quantify soundscape diversity through the lens of functional ecology. 2. Our workflow quantifies the functional diversity of sound in the 24-hour acoustic trait space. We put forward an entity, the Operational Sound Unit (OSU), which groups sounds by their shared functional properties. Using OSUs as our unit of diversity measurement, and building on the framework of Hill numbers, we propose three metrics that capture different aspects of acoustic trait space usage: (i) soundscape richness; (ii) soundscape diversity; (iii) soundscape evenness. We demonstrate the use of these metrics by (a) simulating soundscapes to assess if the indices possess a set of desirable behaviours; and (b) quantifying the soundscape richness and evenness along a gradient in species richness to illustrate how these metrics can be used to shed unique insights into patterns of acoustic niche usage. 3. We demonstrate that: (a) the indices outlined herein have desirable behaviours; and (b) the soundscape richness and evenness are positively correlated with the richness of soniferous species. This suggests that the acoustic niche space is more filled where taxonomic richness is higher. Moreover, species-poor acoustic communities have a higher proportion of rare sounds and use the acoustic space less effectively. As the correlation between the soundscape and taxonomic richness is strong (>0.8) and holds at low sampling intensities, soundscape richness could serve as a proxy for taxonomic richness. 4. Quantifying the soundscape diversity through the lens of functional ecology using the analytical framework of Hill numbers generates novel insights into acoustic niche usage at a landscape scale and provides a useful proxy for taxonomic richness measurement.

Vocal Communication Within a Forest Bird Community

<p><b>This thesis takes a community approach to investigate the acoustics of forest birds in Zealandia sanctuary, Wellington. Initially, the annual changes in vocalisation output of 16 study species and their possible effect on bird conspicuousness were described. Environmental factors that may shape these avian vocalisations were addressed though invoking two key hypotheses, the acoustic adaptation hypothesis, and the acoustic niche hypothesis. In addition, the songs of selected species are investigated: the role of song harmonics in the native North Island saddleback, Philesturnus rufusater, and their role in ranging, change in song dialect through time and space in the introduced song thrush, Turdus philomelos, and temporal change in the song of the native grey warbler, Greygone igata. Vocal activity of the study species varied seasonally, affecting their detectability in bird counts. Some species were mostly first heard rather than seen and viceversa. The results lend support to the acoustic niche hypothesis in that vocalisations within the forest bird community appear to have evolved towards divergence, with native species’ vocalisations being more dispersed within the community acoustic space than those of the introduced species. However, all species concentrated their energy within relatively narrow frequency bands, supporting the predictions of the acoustic adaptation hypothesis. Adaptation to different transmission properties associated with different singing elevations or physiological parameters such as body weight may have an effect on shaping such bird vocalisations.</b></p> <p>Forests provide a complex acoustic space for sound transmission and a “sound window” may not be a constant property of a given forest. The study revealed that a prominent sound window persists in the lower frequency range that is less affected by habitat. Some high frequencies may have similar average attenuation values to those of low frequency, however, with greater fluctuation in attenuation. Ground effect is a further factor in determining how well different frequency ranges transmit and birds may use acoustic characteristics of their habitat to enhance their signal.</p> <p>Harmonics in North Island saddleback chatter song were found to play a potential role in ranging (estimating the distance of signaller), for playback songs with more relative energy within higher harmonics were evidently perceived as coming from a nearby individual.</p> <p>The repertoire size of the song thrush population studied in Zealandia has apparently evolved to become larger and more varied than the source population of song thrushes in the UK, with more syllables delivered with less repetition than the UK song recordings examined.</p> <p>Over a period of 7 years, syllables in grey warbler song have shifted to a higher frequency, but there was no difference in the temporal structure of the song. Habitat effect, competition on the acoustic signal from reintroduced birds and ambient noise level are considered as possible casual factors.</p>

Vocal Communication Within a Forest Bird Community

<p><b>This thesis takes a community approach to investigate the acoustics of forest birds in Zealandia sanctuary, Wellington. Initially, the annual changes in vocalisation output of 16 study species and their possible effect on bird conspicuousness were described. Environmental factors that may shape these avian vocalisations were addressed though invoking two key hypotheses, the acoustic adaptation hypothesis, and the acoustic niche hypothesis. In addition, the songs of selected species are investigated: the role of song harmonics in the native North Island saddleback, Philesturnus rufusater, and their role in ranging, change in song dialect through time and space in the introduced song thrush, Turdus philomelos, and temporal change in the song of the native grey warbler, Greygone igata. Vocal activity of the study species varied seasonally, affecting their detectability in bird counts. Some species were mostly first heard rather than seen and viceversa. The results lend support to the acoustic niche hypothesis in that vocalisations within the forest bird community appear to have evolved towards divergence, with native species’ vocalisations being more dispersed within the community acoustic space than those of the introduced species. However, all species concentrated their energy within relatively narrow frequency bands, supporting the predictions of the acoustic adaptation hypothesis. Adaptation to different transmission properties associated with different singing elevations or physiological parameters such as body weight may have an effect on shaping such bird vocalisations.</b></p> <p>Forests provide a complex acoustic space for sound transmission and a “sound window” may not be a constant property of a given forest. The study revealed that a prominent sound window persists in the lower frequency range that is less affected by habitat. Some high frequencies may have similar average attenuation values to those of low frequency, however, with greater fluctuation in attenuation. Ground effect is a further factor in determining how well different frequency ranges transmit and birds may use acoustic characteristics of their habitat to enhance their signal.</p> <p>Harmonics in North Island saddleback chatter song were found to play a potential role in ranging (estimating the distance of signaller), for playback songs with more relative energy within higher harmonics were evidently perceived as coming from a nearby individual.</p> <p>The repertoire size of the song thrush population studied in Zealandia has apparently evolved to become larger and more varied than the source population of song thrushes in the UK, with more syllables delivered with less repetition than the UK song recordings examined.</p> <p>Over a period of 7 years, syllables in grey warbler song have shifted to a higher frequency, but there was no difference in the temporal structure of the song. Habitat effect, competition on the acoustic signal from reintroduced birds and ambient noise level are considered as possible casual factors.</p>

Timing Is Everything: Acoustic Niche Partitioning in Two Tropical Wet Forest Bird Communities

When acoustic signals sent from individuals overlap in frequency and time, acoustic interference and signal masking may occur. Under the acoustic niche hypothesis (ANH), signaling behavior has evolved to partition acoustic space and minimize overlap with other calling individuals through selection on signal structure and/or the sender’s ability to adjust the timing of signals. Alternately, under the acoustic clustering hypothesis, there is potential benefit to convergence and synchronization of the structural or temporal characteristics of signals in the avian community, and organisms produce signals that overlap more than would be expected by chance. Interactive communication networks may also occur, where species living together are more likely to have songs with convergent spectral and or temporal characteristics. In this study, we examine the fine-scale use of acoustic space in montane tropical wet forest bird communities in Costa Rica and Hawai‘i. At multiple recording stations in each community, we identified the species associated with each recorded signal, measured observed signal overlap, and used null models to generate random distributions of expected signal overlap. We then compared observed vs. expected signal overlap to test predictions of the acoustic niche and acoustic clustering hypotheses. We found a high degree of overlap in the signal characteristics (frequency range) of species in both Costa Rica and Hawai‘i, however, as predicted under ANH, species significantly reduced observed overlap relative to the random distribution through temporal partitioning. There was little support for acoustic clustering or the prediction of the network hypothesis that species segregate across the landscape based on the frequency range of their vocalizations. These findings constitute strong support that there is competition for acoustic space in these signaling communities, and this has resulted primarily in temporal partitioning of the soundscape.

Phantom rivers filter birds and bats by acoustic niche

Natural sensory environments, despite strong potential for structuring systems, have been neglected in ecological theory. Here, we test the hypothesis that intense natural acoustic environments shape animal distributions and behavior by broadcasting whitewater river noise in montane riparian zones for two summers. Additionally, we use spectrally-altered river noise to explicitly test the effects of masking as a mechanism driving patterns. Using data from abundance and activity surveys across 60 locations, over two full breeding seasons, we find that both birds and bats avoid areas with high sound levels, while birds avoid frequencies that overlap with birdsong, and bats avoid higher frequencies more generally. We place 720 clay caterpillars in willows, and find that intense sound levels decrease foraging behavior in birds. For bats, we deploy foraging tests across 144 nights, consisting of robotic insect-wing mimics, and speakers broadcasting bat prey sounds, and find that bats appear to switch hunting strategies from passive listening to aerial hawking as sound levels increase. Natural acoustic environments are an underappreciated niche axis, a conclusion that serves to escalate the urgency of mitigating human-created noise.

Bat echolocation plasticity in allopatry: a call for caution in acoustic identification of Pipistrellus sp.

Animals modify their behaviours and interactions in response to changing environments. In bats, environmental adaptations are reflected in echolocation signalling that is used for navigation, foraging and communication. However, the extent and drivers of echolocation plasticity are not fully understood, hindering our identification of bat species with ultrasonic detectors, particularly for cryptic species with similar echolocation calls. We used a combination of DNA barcoding, intensive trapping, roost and emergence surveys and acoustic recording to study a widespread European cryptic species complex (Pipistrellus pipistrellus and Pipistrellus pygmaeus) to investigate whether sibling bat species could exhibit extreme echolocation plasticity in response to certain environmental conditions or behaviours. We found that P. pygmaeus occupied the acoustic niche of their absent congeneric species, producing calls with P. pipistrellus’ characteristic structure and peak frequencies and resulting in false positive acoustic records of that species. Echolocation frequency was significantly affected by the density of bats and by maternity rearing stage, with lower frequency calls emitted when there was a high density of flying bats, and by mothers while juveniles were non-volant. During roost emergence, 29% of calls had peak frequencies typical of P. pipistrellus, with calls as low as 44 kHz, lower than ever documented. We show that automatic and manual call classifiers fail to account for echolocation plasticity, misidentifying P. pygmaeus as P. pipistrellus. Our study raises a vital limitation of using only acoustic sampling in areas with high densities of a single species of a cryptic species pair, with important implications for bat monitoring. Significance statement Ultrasonic acoustic detectors are widely used in bat research to establish species inventories and monitor species activity through identification of echolocation calls, enabling new methods to study and understand this elusive understudied group of nocturnal mammals. However, echolocation call signalling in bats is intrinsically different to that of other taxa, serving a main function of navigation and foraging. This study demonstrates an extreme level of plasticity, showing large variation in call frequency and structure in different situations. We showcase the difficulty and limitation in using acoustic sampling alone for bat monitoring and the complications of setting parameters for species identification for manual and automatic call classifiers. Our observations of call frequency variation correlated with density and absence of congenerics provide novel insights of behavioural echolocation plasticity in bats.

O NICHO ACÚSTICO: INTEGRANDO A FÍSICA, ECOLOGIA E TEORIA DA COMUNICAÇÃO

Animal communication is related to many biological processes, and its investigation may help elucidate many aspects of an organism. If in the one hand, animal communication might be seen as a biological process, in the other, sound is a physical phenomenon, so that the understanding of animal communication depends on processes that go beyond biology itself. Here we aim to interlace concepts of Physics, Ecology, and communication to better operationalize the concept of acoustic niche. The study of the relationships between the communication problems (i.e. transmission, semantics and effectiveness), the physical properties of sound (frequency, intensity and time), and ecological aspects of communication, permits a better understanding of the acoustic niche and its evolution.

Acoustic space partition by four Hylidae in an open habitat in Central Brazil

The use and partition of the acoustic space by anuran species is of special interest for evaluating inter-specific interactions in acoustic traits. Here we evaluate the acoustic space partitioning by four species of hylidae over the course of eleven months, at the Ecological Station of Águas Emendadas - DF, central Brazil. The acoustic niche overlap among species was smaller than expected by chance, indicating a competition in the use of this resource. If competition affects the use of acoustic resouces by species, low overlap on advertisement call traits are relevant in open habitats anuran assemblies.