Puppy whines mediate maternal behavior in domestic dogs.

In mammals, offspring vocalizations typically encode information about identity and body condition, allowing parents to limit alloparenting and adjust care. But how do these vocalizations mediate parental behavior in species faced with the problem of rearing not one, but multiple offspring, such as domestic dogs? Comprehensive acoustic analyses of 4,400 whines recorded from 220 Beagle puppies in 40 litters revealed litter and individual (within litter) differences in call acoustic structure. By then playing resynthesized whines to mothers, we showed that they provided more care to their litters, and were more likely to carry the emitting loudspeaker to the nest, in response to whine variants derived from their own puppies than from strangers. Importantly, care provisioning was attenuated by experimentally moving the fundamental frequency (fo, perceived as pitch) of their own puppies' whines outside their litter-specific range. Within most litters, we found a negative relationship between puppies' whine fo and body weight. Consistent with this, playbacks showed that maternal care was stronger in response to high-pitched whine variants simulating relatively small offspring within their own litter's range compared to lower-pitched variants simulating larger offspring. We thus show that maternal care in a litter-rearing species relies on a dual assessment of offspring identity and condition, largely based on level-specific inter- and intra-litter variation in offspring call fo. This dual encoding system highlights how, even in a long-domesticated species, vocalizations reflect selective pressures to meet species-specific needs. Comparative work should now investigate whether similar communication systems have convergently evolved in other litter-rearing species.

Beyond speech: Exploring diversity in the human voice.

Humans have evolved voluntary control over vocal production for speaking and singing, while preserving the phylogenetically older system of spontaneous nonverbal vocalizations such as laughs and screams. To test for systematic acoustic differences between these vocal domains, we analyzed a broad, cross-cultural corpus representing over 2 h of speech, singing, and nonverbal vocalizations. We show that, while speech is relatively low-pitched and tonal with mostly regular phonation, singing and especially nonverbal vocalizations vary enormously in pitch and often display harsh-sounding, irregular phonation owing to nonlinear phenomena. The evolution of complex supralaryngeal articulatory spectro-temporal modulation has been critical for speech, yet has not significantly constrained laryngeal source modulation. In contrast, articulation is very limited in nonverbal vocalizations, which predominantly contain minimally articulated open vowels and rapid temporal modulation in the roughness range. We infer that vocal source modulation works best for conveying affect, while vocal filter modulation mainly facilitates semantic communication.

Nonlinear vocal phenomena affect human perceptions of distress, size and dominance in puppy whines.

While nonlinear phenomena (NLP) are widely reported in animal vocalizations, often causing perceptual harshness and roughness, their communicative function remains debated. Several hypotheses have been put forward: attention-grabbing, communication of distress, exaggeration of body size and dominance. Here, we use state-of-the-art sound synthesis to investigate how NLP affect the perception of puppy whines by human listeners. Listeners assessed the distress, size or dominance conveyed by synthetic puppy whines with manipulated NLP, including frequency jumps and varying proportions of subharmonics, sidebands and deterministic chaos. We found that the presence of chaos increased the puppy's perceived level of distress and that this effect held across a range of representative fundamental frequency (fo) levels. Adding sidebands and subharmonics also increased perceived distress among listeners who have extensive caregiving experience with pre-weaned puppies (e.g. breeders, veterinarians). Finally, we found that whines with added chaos, subharmonics or sidebands were associated with larger and more dominant puppies, although these biases were attenuated in experienced caregivers. Together, our results show that nonlinear phenomena in puppy whines can convey rich information to human listeners and therefore may be crucial for offspring survival during breeding of a domesticated species.

Friend or foe: Risso's dolphins eavesdrop on conspecific sounds to induce or avoid intra-specific interaction.

The detection and use of emitters' signals by unintended receivers, i.e., eavesdropping, represents an important and often low-cost way for animals to gather information from their environment. Acoustic eavesdropping can be a key driver in mediating intra- and interspecific interactions (e.g., cooperation, predator-prey systems), specifically in species such as cetaceans that use sound as a primary sensory modality. While most cetacean species produce context-specific sounds, little is known about the use of those sounds by potential conspecific eavesdroppers. We experimentally tested the hypothesis that a social cetacean, Risso's dolphin (Grampus griseus), is able to gather biologically relevant information by eavesdropping on conspecific sounds. We conducted playback experiments on free-ranging dolphins using three context-specific sounds stimuli and monitored their horizontal movement using visual or airborne focal follow observations. We broadcasted natural sequences of conspecific foraging sounds potentially providing an attractive dinner bell signal (n = 7), male social sounds simulating a risk of forthcoming agonistic interaction (n = 7) and female-calf social sounds representing no particularly threatening context (n = 7). We developed a quantitative movement response score and tested whether animals changed their direction of horizontal movement towards or away from the playback source. Dolphins approached the foraging and the social female-calf sounds whereas they avoided the social male sounds. Hence, by acoustically eavesdropping on conspecifics, dolphins can discriminate between social and behavioural contexts and anticipate potential threatening or beneficial situations. Eavesdropping and the ensuing classification of 'friend or foe' can thus shape intra-specific social interactions in cetaceans.

Harsh is large: nonlinear vocal phenomena lower voice pitch and exaggerate body size.

A lion's roar, a dog's bark, an angry yell in a pub brawl: what do these vocalizations have in common? They all sound harsh due to nonlinear vocal phenomena (NLP)-deviations from regular voice production, hypothesized to lower perceived voice pitch and thereby exaggerate the apparent body size of the vocalizer. To test this yet uncorroborated hypothesis, we synthesized human nonverbal vocalizations, such as roars, groans and screams, with and without NLP (amplitude modulation, subharmonics and chaos). We then measured their effects on nearly 700 listeners' perceptions of three psychoacoustic (pitch, timbre, roughness) and three ecological (body size, formidability, aggression) characteristics. In an explicit rating task, all NLP lowered perceived voice pitch, increased voice darkness and roughness, and caused vocalizers to sound larger, more formidable and more aggressive. Key results were replicated in an implicit associations test, suggesting that the 'harsh is large' bias will arise in ecologically relevant confrontational contexts that involve a rapid, and largely implicit, evaluation of the opponent's size. In sum, nonlinearities in human vocalizations can flexibly communicate both formidability and intention to attack, suggesting they are not a mere byproduct of loud vocalizing, but rather an informative acoustic signal well suited for intimidating potential opponents.

Evidence for discrimination between feeding sounds of familiar fish and unfamiliar mammal-eating killer whale ecotypes by long-finned pilot whales.

Killer whales (KW) may be predators or competitors of other cetaceans. Since their foraging behavior and acoustics differ among populations ('ecotypes'), we hypothesized that other cetaceans can eavesdrop on KW sounds and adjust their behavior according to the KW ecotype. We performed playback experiments on long-finned pilot whales (Globicephala melas) in Norway using familiar fish-eating KW sounds (fKW) simulating a sympatric population that might compete for foraging areas, unfamiliar mammal-eating KW sounds (mKW) simulating a potential predator threat, and two control sounds. We assessed behavioral responses using animal-borne multi-sensor tags and surface visual observations. Pilot whales barely changed behavior to a broadband noise (CTRL-), whereas they were attracted and exhibited spyhops to fKW, mKW, and to a repeated-tonal upsweep signal (CTRL+). Whales never stopped nor started feeding in response to fKW, whereas they reduced or stopped foraging to mKW and CTRL+. Moreover, pilot whales joined other subgroups in response to fKW and CTRL+, whereas they tightened individual spacing within group and reduced time at surface in response to mKW. Typical active intimidation behavior displayed to fKW might be an antipredator strategy to a known low-risk ecotype or alternatively a way of securing the habitat exploited by a heterospecific sympatric population. Cessation of feeding and more cohesive approach to mKW playbacks might reflect an antipredator behavior towards an unknown KW ecotype of potentially higher risk. We conclude that pilot whales are able to acoustically discriminate between familiar and unfamiliar KW ecotypes, enabling them to adjust their behavior according to the perceived disturbance type.