Ole Næsbye Larsen

The Source-Path-Receiver Model (SPRM) is a fundamental concept derived from hazard (including noise) control. It is useful in studies of animal bioacoustics where the sound sources may be animals, humans, or natural events within the habitat and the receivers are animals. It provides a framework for the researcher to ensure all aspects of the scenario being observed or recorded are considered, which could affect the observations. This chapter develops the SPRM for the example of animal acoustic communication, where the source and receiver are animals of the same species. Factors that affect the source and receiver are explained (e.g., age, sex, individual identity, and context). Much emphasis is given to “the path.” The environment through which the sound travels affects the received signal (in terms of its amplitude, frequency, and duration) and exhibits ambient noise, which might affect both sender and receiver. The basic concepts of sound propagation are explained (including Huygens’ principle, ray tracing, Snell’s law, reflection, scattering, reverberation, diffraction, refraction, transmission, absorption, ground effect, atmosphere effects, acoustic mirages, and shadow zones). The SPRM illustrates the importance of exploring the acoustic features of a sound signal at all points between the sender and receiver to understand factors that could promote or inhibit effective communication among animals.

Human speech production obeys the same acoustic principles as vocal production in other animals but has distinctive features: A stable vocal source is filtered by rapidly changing formant frequencies. To understand speech evolution, we examined a wide range of primates, combining observations of phonation with mathematical modeling. We found that source stability relies upon simplifications in laryngeal anatomy, specifically the loss of air sacs and vocal membranes. We conclude that the evolutionary loss of vocal membranes allows human speech to mostly avoid the spontaneous nonlinear phenomena and acoustic chaos common in other primate vocalizations. This loss allows our larynx to produce stable, harmonic-rich phonation, ideally highlighting formant changes that convey most phonetic information. Paradoxically, the increased complexity of human spoken language thus followed simplification of our laryngeal anatomy.

When suitable nesting habitat is rare, birds may have to share it with heterospecific individuals with similar nesting requirements. The resulting species mosaic may in turn affect how breeding birds communicate vocally with each other. For instance, they may modify their vocalizations if the ambient noise produced by heterospecifics interferes with their own vocalizations. The colonies of Gentoo Penguins (Pygoscelis papua ellsworthi) are interesting to examine whether such a vocal variation occurs in colonial seabirds, as these birds frequently breed in the same colony as Adélie (P. adeliae) or Chinstrap (P. antarcticus) penguins. We investigated whether the presence of congeneric penguins breeding in the same colony as Gentoo Penguins affected the acoustic characteristics of their most common vocalization, the ecstatic display call. Based on vocalizations recorded in 23 breeding colonies along the Antarctic Peninsula during 3 field seasons, we found that the frequency of ecstatic display calls of Gentoo Penguins was consistently lower (average exhale phrase ~460 Hz lower, average inhale phrase ~370 Hz lower) as well as decreased in energy distribution when they bred in mixed colonies with Adélie Penguins than when they bred only in proximity to conspecifics. In contrast, the frequency of both inhale and exhale phrases was unaffected by the presence of Chinstrap Penguins, potentially due to the already greater frequency difference between these 2 species. The apparent vocal plasticity, in mixed colonies with Adélie Penguins, likely allows Gentoo Penguins to adjust the quality of their vocalizations depending on the acoustic space available in their colonies, and hence possibly enhance signal transmission between conspecifics. However, whether this acoustic adjustment is sufficient to increase the detectability of vocalizations and localization of individuals has yet to be determined. • We examined whether Gentoo Penguin calls differ between breeding colonies that included multiple species of penguins and those where only Gentoo Penguins bred. • We compared call parameters including frequency, structure, and duration between individual Gentoo Penguins breeding in colonies with and without the presence of Adélie or Chinstrap penguins along the Antarctic Peninsula during three austral summers. • We found that the calls of Gentoo Penguins showed a lower frequency and narrower energy distribution when they bred in proximity to Adélie Penguins but did not change when they bred in proximity to Chinstrap Penguins with higher main frequency vocalizations.. • This suggests that penguins can adjust the quality of their vocalizations according to the social (and presumably acoustic) environment of their colonies.

Male walruses produce some of the longest continuous reproductive displays known among mammals to convey their physical fitness to potential rivals and possibly to potential mates. Here, we document the ability of a captive walrus to produce intense, rhythmic sounds through a non-vocal pathway involving deliberate, regular collision of the fore flippers. High-speed videography linked to an acoustic onset marker revealed sound production through cavitation, with the acoustic impulse generated by each forceful clap exceeding a peak-to-peak sound level of 200 dB re. 1 µPa. This clapping display is in some ways quite similar to the knocking display more commonly associated with walruses in rut but is produced through a very different mechanism and with much higher amplitudes. While this clapping behaviour has not yet been documented in wild individuals, it has been observed among other mature male walruses living in human care. Production of intense sounds through cavitation has previously been documented only in crustaceans but may also be an effective means of sound production for some aquatic mammals.

Wildlife species are often host to a diversity of parasites, but our knowledge of their diversity and ecology is extremely limited, especially for reptiles. Little is known about the host-parasite ecology of the Australian lizard, the King's skink (Egernia kingii). In spring of 2015, we carried out a field-based study of a population of King's skinks on Penguin Island (Western Australia). We documented five species of parasites, including two ectoparasitic mites (an undescribed laelapid mite and Mesolaelaps australiensis), an undescribed coccidia species, and two nematode species (Pharyngodon tiliquae and Capillaria sp.). The laelapid mite was the most abundant parasite, infesting 46.9% of the 113 captured lizards. This mite species increased in prevalence and abundance over the course of the study. Infection patterns of both mites varied with lizard life-stage; sub-adults were more commonly infested with laelapid mites than adults or juveniles, and sub-adults and adults were infested by more laelapid mites than juveniles. By contrast, adults had a higher prevalence of M. australiensis than juveniles or sub-adults. Among the gastrointestinal parasites, P. tiliquae was relatively common among the sampled lizards (35.3%). These results give new important information about reptiles as parasite hosts and what factors influence infection patterns.

Hearing is vital for birds as they rely on acoustic communication with parents, mates, chicks, and conspecifics. Amphibious seabirds face many ecological pressures, having to sense cues in air and underwater. Natural noise conditions have helped shape this sensory modality but anthropogenic noise is increasingly impacting seabirds. Surprisingly little is known about their hearing, despite their imperiled status. Understanding sound sensitivity is vital when we seek to manage manmade noise impacts. We measured the auditory sensitivity of nine wild Atlantic puffins, Fratercula arctica, in a capture-and-release setting in an effort to define their audiogram and compare these data to the hearing of other birds and natural rookery noise. Auditory sensitivity was tested using auditory evoked potential (AEP) methods. Responses were detected from 0.5 to 6 kHz. Mean thresholds were below 40 dB re 20 µPa from 0.75 to 3 kHz indicating that these were the most sensitive auditory frequencies, similar to other seabirds. Thresholds in the ‘middle’ frequency range 1-2.5 kHz were often down to 10-20 dB re 20 µPa. Lowest thresholds were typically at 2.5 kHz. These are the first in-air auditory sensitivity data from multiple wild-caught individuals of a deep-diving Alcid seabird. The audiogram was comparable to other birds of similar size, thereby indicating that puffins have fully functioning aerial hearing despite the constraints of their deep-diving, amphibious lifestyles. There was some variation in thresholds, yet animals generally had sensitive ears suggesting aerial hearing is an important sensory modality for this taxon.

Diving birds spend up to several minutes underwater during pursuit-dive foraging. To find and capture prey, like fish and squid, they probably need several senses in addition to vision. Cormorants, very efficient predators of fishes, have unexpectedly low visual acuity underwater. So, underwater hearing may be an important sense, as for other diving animals. We measured auditory thresholds and eardrum vibrations in air and underwater of the great cormorant (Phalacrocorax carbo sinensis). Wild-caught cormorant fledglings were anesthetized, and their auditory brainstem response (ABR) and eardrum vibrations to clicks and tone bursts were measured, first in an anechoic box in air and then in a large water-filled tank, with their head and ears submerged 10 cm below the surface. Both the ABR-response waveshape and latency, as well as the ABR-thresholds, measured in units of sound pressure, were similar in air and water. The best average sound pressure sensitivity was found at 1 kHz, both in air (53 dB re. 20 µPa) and underwater (58 dB re. 20 µPa). When thresholds were compared in units of intensity, however, the sensitivity underwater was higher than in air. Eardrum vibration amplitudes in both media reflected the ABR-threshold curves. These results suggest that cormorants have in-air hearing abilities comparable to similar-sized diving birds, and that their underwater hearing sensitivity is at least as good as their aerial sensitivity. This together with the morphology of the outer ear (collapsible meatus) and middle ear (thickened eardrum), suggest that cormorants may have anatomical and physiological adaptations for amphibious hearing.

Cooperation is a complex behaviour found in many kinds of organisms and occurs between individuals of the same and different species. Several studies have examined the intentionality of this behaviour by testing the animals’ understanding of the need for a partner when working in pairs. The mammalian species tested express such understanding, whereas most tested birds fail, especially when the test involves a delayed access to the setup by one of the co-operators. In the present study, the cooperative problem-solving capability of four peach-fronted conures (Eupsittula aurea) was investigated with the loose string test. All four parrots solved the paradigm by simultaneously pulling the ends of the same string to bring a platform with a food reward within reach. They were also capable of solving the task when one of the co-operators was delayed, even when visually isolated from each other. To further test their comprehension and to exclude the birds relying on task-associated cues, we video-recorded the trials and quantified possible cues and strategies for timing the pulling behaviour (e.g., sound of the partner’s door when opening, sound of steps of partner approaching). The preferred cue to start pulling was to wait for their partner’s arrival to the string. The number of vocalisations was significantly higher during visually isolated conditions and for successful trials compared to failed trials, suggesting possible information exchange. Our findings show that peach-fronted conures can solve a cooperative task, and that cooperation success is not determined by external cues or by partner identity or affinity.