Thierry Lengagne

Finding a parent in a king penguin colony: the acoustic system of individual recognition

To be fed, a king penguin, Aptenodytes patagonicus, chick must identify the call of its parents, in the continuous background noise of the colony. To study this recognition process, we played back to the chicks parental calls with acoustic parameters modified in the temporal and frequency domains. The parental call is composed of syllables (complex sounds with harmonic series) separated by pronounced amplitude declines. Our experiments with modified signals indicate that the chicks frequency analysis of the call is not tuned towards precise peak energy values, the signal being recognized even when the carrier frequency was shifted 100 Hz down or 75 Hz up. To recognize the adult, chicks used frequency rather than amplitude modulation, in particular the frequency modulation shape of the syllable. This structure is repeated through the different syllables of the call giving a distinct vocal signature. Our experiments also show that the receiver needs to perceive only a small part of the signal: the first half of the syllable (0.23 s) and the first three harmonics were sufficient to elicit recognition. The small amount of information necessary to understand the message, the high redundancy in the time and frequency domains and the almost infinite possibilities of coding provided by the frequency modulation signature permit the chick to recognize the adult, without the help of a nest site. For these reasons, the code used in the call of the king penguin can be regarded as a functional code, increasing the possibility of individual recognition in an acoustically constraining environment. Copyright 1999 The Association for the Study of Animal Behaviour.

How do king penguins (Aptenodytes patagonicus apply the mathematical theory of information to communicate in windy conditions

In the king penguin (Aptenodytes patagonicus), both pair members alternate in incubating and rearing their chick. Mates can recognize each other among thousands of other birds in the hubbub of the colony using only acoustic signalling: the display call. Large penguin colonies are found on sub–Antarctic islands where strong winds blow throughout the year. We have shown by experiments under natural conditions that the level of background noise increases in windy conditions and thus leads to a diminution of the signal–to–noise ratio. Moreover the emergence level of the signal revealed by entropy calculation is statistically weaker in windy conditions. To achieve breeding success, birds must continue communicating in spite of the significant decrease in the total amount of information that can be transmitted in windy situations. For the first time, to our knowledge, we have shown that a bird species takes into account the constraints imposed by wind on their acoustic communication. In windy conditions, birds try to maintain the efficiency of communication by increasing both the number of calls emitted and the number of syllables per call. This result conforms with predictions from the mathematical theory of communication: increased redundancy in a signal improves the probability of receiving a message in a noisy channel.

The effects of rain on acoustic communication: tawny owls have good reason for calling less in wet weather.

Numerous attempts have been made to quantify ecological factors that affect the calling range of animal signals. The various processes leading signals to become distorted and embedded in background noise have been described in many habitats (ranging from forest to savannah) and the propagation path in these biomes has thereby been characterized. However, the impact of climatic factors on acoustic communication has been little studied. Surprisingly, to our knowledge, the importance of rain, a regular phenomenon occurring in all habitats except deserts, has never been investigated. Here, we describe a 69–fold advantage in area reached by the call of a territorial bird, the tawny owl (Strix aluco) in dry versus rainy conditions. In support of this, we found a marked reduction in the calling of tawny owls in rainy conditions. Constraints imposed by a rainy propagation path are likely to modify the reliability of acoustic information and thus calling behaviour of many animals.

Genetic bottlenecks driven by population disconnection.

Connectivity among populations plays a crucial role in maintaining genetic variation at a local scale, especially in small populations affected strongly by genetic drift. The negative consequences of population disconnection on allelic richness and gene diversity (heterozygosity) are well recognized and empirically established. It is not well recognized, however, that a sudden drop in local effective population size induced by such disconnection produces a temporary disequilibrium in allelic frequency distributions that is akin to the genetic signature of a demographic bottleneck. To document this effect, we used individual-based simulations and empirical data on allelic richness and gene diversity in six pairs of isolated versus well-connected (core) populations of European tree frogs. In our simulations, population disconnection depressed allelic richness more than heterozygosity and thus resulted in a temporary excess in gene diversity relative to mutation drift equilibrium (i.e., signature of a genetic bottleneck). We observed a similar excess in gene diversity in isolated populations of tree frogs. Our results show that population disconnection can create a genetic bottleneck in the absence of demographic collapse.

Multiple signals and male spacing affect female preference at cocktail parties in treefrogs

Effective acoustic communication in the face of intense conspecific background noise constitutes a constant sensory challenge in chorusing and colonial species. An evolutionary approach suggests that behavioural and environmental constraints in these species should have shaped signal design and signalling behaviour to enable communication in noisy conditions. This could be attained both through the use of multicomponent signals and through short-term adjustments in the spatial separation of calling males. We investigated these two hypotheses in a chorusing anuran, the hylid Hyla arborea, through a series of phonotaxis experiments conducted within a six-speaker arena in a high background noise situation, by presenting females with male calls containing either single or multiple attractive call components, and by modifying distances between speakers. We found that female ability to discriminate attractive calls increased when several attractive call components were available, providing novel evidence that the use of multicomponent signals enhances communication in complex acoustic conditions. Signal discrimination in females also improved with speaker separation, demonstrating that within natural choruses, spatial unmasking conditioned by male density and spatial separation probably improves female discrimination of competing males. Implications of these results for the accuracy of mate choice within choruses are discussed.

A method of independent time and frequency decomposition of bioacoustic signals: inter-individual recognition in four species of penguins

The aim of this paper is to introduce a method for analyzing acoustic signals capable of assessing the potential for individual coding information. Signals are analysed both in the time domain (rhythm of emission of the song independent of its frequency content) and in the spectral domain (spectral content of the song independent of the rhythm of emission). The method is then applied to a comparative study of four penguin species, where the problem posed by inter-individual recognition differs from species to species. A direct relationship was shown between the potential of individual coding and the difficulty in partner identification.

GENETIC EROSION IN WILD POPULATIONS MAKES RESISTANCE TO A PATHOGEN MORE COSTLY

Populations that have suffered from genetic erosion are expected to exhibit reduced average trait values or decreased variation in adaptive traits when experiencing periodic or emergent stressors such as infectious disease. Genetic erosion may consequentially modify the ability of a potential host population to cope with infectious disease emergence. We experimentally investigate this relationship between genetic variability and host response to exposure to an infectious agent both in terms of susceptibility to infection and indirect parasite‐mediated responses that also impact fitness. We hypothesized that the deleterious consequences of exposure to the pathogen (Batrachochytrium dendrobatidis) would be more severe for tadpoles descended from European treefrog (Hyla arborea) populations lacking genetic variability. Although all exposed tadpoles lacked detectable infection, we detected this relationship for some indirect host responses, predominantly in genetically depleted animals, as well as an interaction between genetic variability and pathogen dose on life span during the postmetamorphic period. Lack of infection and a decreased mass and postmetamorphic life span in low genetic diversity tadpoles lead us to conclude that genetic erosion, while not affecting the ability to mount effective resistance strategies, also erodes the capacity to invest in resistance, increased tadpole growth rate, and metamorphosis relatively simultaneously.

The challenge of finding a high-quality male: a treefrog solution based on female assessment of male calls

In the context of sexual selection through mate choice, it has recently been suggested that the effectiveness of intersexual communication is affected not only by the honesty of male sexual signals but also by the number of signal components simultaneously used to convey information. We investigated female use of acoustic components in the chorusing treefrog Hyla arborea using two-choice phonotaxis tests. Females showed a significant preference for higher call rates and calls of short duration, as well as for higher call amplitudes. They also favoured lower peak frequencies. Since this call characteristic was negatively correlated with body weight, such a preference should lead to their selecting larger males. Females also exhibited a marginal preference for longer call bouts. The reliability of the multiple call components involved in mate choice in H. arborea are discussed, as well as the validity of these results for mate choice in the natural context of a noisy anuran chorus.

Male mating speed promote hybridization in the Rana lessonae–Rana esculenta waterfrog system

Hybridization is a widespread phenomenon in many vertebrate groups. Prezygotic isolating mechanisms, probably caused by selection against hybrids with reduced fitness, reduce the likelihood of such events. Although hybrid-reduced fitness relatively to parental species is common, hybridization can also be beneficial, and hybrids sometimes outperform the pure species type. In this study, we examined two potential processes, Hubbs’s principle and male–male competition, which could enhance hybridization in the waterfrog complex and thus explain the proportion of heterospecific pairs collected in a natural pond. Firstly, by collecting 791 frogs in the field to study pair and chorus composition, we showed that in a mixed Rana lessonae–Rana esculenta population, the scarcity of hybrid R. esculenta males did not account for the proportion of heterospecific pairs: indeed, when examining pairing composition in six different choruses, we found that hybrid males were always under-represented and that R. esculenta females were found paired with R. lessonae males. Secondly, we investigated experimentally whether or nor male–male competition mechanism could explain pair formation in waterfrogs. Our mating speed experiment highlights mechanisms that could explain heterospecific pairs in a context of promiscuous mating where scramble competition was intense. To measure the rapidity with which a male grasps a female, we placed males in a grid cage with a female, and the dynamics of pair formation was monitored. R. esculenta males showed a lower pairing success than R. lessonae males as a smaller proportion of them amplexed females, and more time was needed for them to get amplexed. Thus, a less adaptative mechanism than female mate choice may also explain the mating pattern observed in waterfrog species.

Consequences of genetic erosion on fitness and phenotypic plasticity in European tree frog populations (Hyla arborea)

The detrimental effects of genetic erosion on small isolated populations are widely recognized contrary to their interactions with environmental changes. The ability of genotypes to plastically respond to variability is probably essential for the persistence of these populations. Genetic erosion impact may be exacerbated if inbreeding affects plastic responses or if their maintenance were at higher phenotypic costs. To understand the interplay ‘genetic erosion–fitness–phenotypic plasticity’, we experimentally compared, in different environments, the larval performances and plastic responses to predation of European tree frogs (Hyla arborea) from isolated and connected populations. Tadpoles from isolated populations were less performant, but the traits affected were environmental dependant. Heterosis observed in crosses between isolated populations allowed attributing their low fitness to inbreeding. Phenotypic plasticity can be maintained in the face of genetic erosion as inducible defences in response to predator were identical in all populations. However, the higher survival and developmental costs for isolated populations in harsh conditions may lead to an additional fitness loss for isolated populations.