Crocodilians are among the most vocal non-avian reptiles. Adults of both sexes produce loud vocalizations known as ‘bellows’ year round, with the highest rate during the mating season. Although the specific function of these vocalizations remains unclear, they may advertise the caller's body size, because relative size differences strongly affect courtship and territorial behaviour in crocodilians. In mammals and birds, a common mechanism for producing honest acoustic signals of body size is via formant frequencies (vocal tract resonances). To our knowledge, formants have to date never been documented in any non-avian reptile, and formants do not seem to play a role in the vocalizations of anurans. We tested for formants in crocodilian vocalizations by using playbacks to induce a female Chinese alligator (Alligator sinensis) to bellow in an airtight chamber. During vocalizations, the animal inhaled either normal air or a helium/oxygen mixture (heliox) in which the velocity of sound is increased. Although heliox allows normal respiration, it alters the formant distribution of the sound spectrum. An acoustic analysis of the calls showed that the source signal components remained constant under both conditions, but an upward shift of high-energy frequency bands was observed in heliox. We conclude that these frequency bands represent formants. We suggest that crocodilian vocalizations could thus provide an acoustic indication of body size via formants. Because birds and crocodilians share a common ancestor with all dinosaurs, a better understanding of their vocal production systems may also provide insight into the communication of extinct Archosaurians.
In many vertebrates, acoustic cues to body size are encoded in resonance frequencies of the vocal tract (“formants”), rather than in the rate of tissue vibration in the sound source (“pitch”). Anatomical constraints on the vocal tract’s size render formants honest cues to size in many bird and mammal species, but it is not clear whether this correlation evolved convergently in these two clades, or whether it is widespread among amniotes (mammals, birds, and non-avian reptiles). We investigated the potential for honest acoustic cues in the bellows of adult American alligators and found that formant spacing provided highly reliable cues to body size, while presumed correlates of the source signal did not. These findings held true for both sexes and for all bellows whether produced in or out of water. Because birds and crocodilians are the last extant Archosaurians and share common ancestry with all extinct dinosaurs, our findings support the hypothesis that dinosaurs used formants as cues to body size. The description of formants as honest signals in a non-avian reptile combined with previous evidence from birds and mammals strongly suggests that the principle of honest signalling via vocal tract resonances may be a broadly shared trait among amniotes.
Behavioral predispositions are innate tendencies of animals to behave in a given way without the input of learning. They increase survival chances and, due to environmental and ecological challenges, may vary substantially even between closely related taxa. These differences are likely to be especially pronounced in long-lived species like crocodilians. This order is particularly relevant for comparative cognition due to its phylogenetic proximity to birds. Here we compared early life behavioral predispositions in two Alligatoridae species. We exposed American alligator and spectacled caiman hatchlings to three different novel situations: a novel object, a novel environment that was open and a novel environment with a shelter. This was then repeated a week later. During exposure to the novel environments, alligators moved around more and explored a larger range of the arena than the caimans. When exposed to the novel object, the alligators reduced the mean distance to the novel object in the second phase, while the caimans further increased it, indicating diametrically opposite ontogenetic development in behavioral predispositions. Although all crocodilian hatchlings face comparable challenges, e.g., high predation pressure, the effectiveness of parental protection might explain the observed pattern. American alligators are apex predators capable of protecting their offspring against most dangers, whereas adult spectacled caimans are frequently predated themselves. Their distancing behavior might be related to increased predator avoidance and also explain the success of invasive spectacled caimans in the natural habitats of other crocodilians.
Human language involves combining items into meaningful, syntactically structured wholes. The evolutionary origin of syntactic abilities has been investigated by testing pattern perception capacities in nonhuman animals. New World primates can respond spontaneously to structural changes in acoustic sequences and songbirds can learn to discriminate between various patterns in operant tasks. However, there is no conclusive evidence that songbirds respond spontaneously to structural changes in patterns without reinforcement or training. In this study, we tested pattern perception capacities of common ravens, Corvus corax, in a habituation–discrimination playback experiment. To enhance stimulus salience, call recordings of male and female ravens were used as acoustic elements, combined to create artificial territorial displays as target patterns. We habituated captive territorial raven pairs to displays following a particular pattern and subsequently exposed them to several test and control playbacks. Subjects spent more time visually orienting towards the loudspeaker in the discrimination phase when they heard structurally novel call combinations, violating the pattern presented during habituation. This demonstrates that songbirds, much like primates, can be sensitive to structural changes in auditory patterns and respond to them spontaneously, without training.
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