Abstract: Lacertid lizards have been hailed as a model system for the study of reptilian chemical communication. However, results obtained with the genus Podarcis, a diverse group of wall lizards with complex systematics, challenge emerging paradigms and caution against hasty generalizations. Here we review the available evidence on the role of chemical stimuli in male-female and male-male interactions in Iberian Podarcis. Males of several species can discriminate between chemicals left on substrates by females of their own or a different species, suggesting that differences in female chemical cues may underlie species recognition in this group. Females, on the other hand, do not respond differentially to conspecific and congeneric male scent marks. Males of Podarcis liolepis use scent marks to recognize rivals individually, evaluate their competitive ability (i.e., body size), and assess the threat posed by each individual rival neighbor. In contrast, females do not exhibit a preference for territories scent marked by larger (i.e., more competitive) males, which suggests a limited role for male scent marks in pre-copulatory mate choice. This behavioral sex difference is consistent with detailed neuro-ethological evidence showing that chemosensory brain areas in P. liolepis are sexually dimorphic. The accessory olfactory bulbs are larger (both in absolute and relative terms) in males than in females, probably as a result of sex-specific rates of adult neurogenesis. In both sexes, cell proliferation undergoes seasonal cycles that may have evolved to satisfy increased chemosensory demands at particular times of the year. Overall, and against recent generalizations, these results suggest that male scent marks have been shaped mainly by strong intrasexual selection.
Abstract: Under certain circumstances, prey may inform potential predators of their unprofitability by means of pursuit-deterrent signals. The evidence for pursuit-deterrent signaling in reptiles is scant and taxonomically biased. Wall lizards, Podarcis muralis (Squamata: Lacertidae) produce several distinct types of stereotyped foot shake displays, of which one, performed in antipredator contexts, is a likely candidate for a pursuit-deterrent function. We investigated this possibility by recording the responses of lizards in the field to a slowly approaching human acting as a surrogate predator. In addition to starting and flight initiation distances, we measured the presence of foot shakes, the leg that was shaken, and the distance from the observer at which the display was performed (display distance). Of a total of 484 approaches, 109 (22.5%) elicited foot shake displays. Roughly half the lizards displayed from the location where they were first sighted, while the other half moved a short distance, then displayed. There was no left-right preference in the leg used to display, but most lizards displayed with the leg closer to the approaching predator. Juveniles and subadults had smaller flight initiation distances than adult lizards. There were no sex-related differences in starting or flight initiation distances, but females, for a given distance, were more likely to display than males. Foot shake display frequency declined abruptly at 1 m. If lizards waited until the surrogate predator was this close, they mostly fled without displaying. Our results show that antipredator foot shaking in P. muralis is consistent with expectations from pursuit-deterrent theory.
Abstract: Female mate choice based on male phenotypic traits is controversial in lizards, particularly in territorial species. In this study, we examine female choice of male scent marks in a territorial lacertid lizard (Podarcis hispanica) in which scent marks have been shown to signal male size (i.e., an important determinant of competitive ability in this species). Females were simultaneously exposed to three naturalized 4 m² choice areas bearing: (1) no scent marks (i.e., control), (2) scent marks of large males, and (3) scent marks of small males. Although female lizards preferentially associated with scent marked choice areas, we found no evidence that females chose territories marked by large males. Furthermore, in response to experimentally induced dusk at the end of choice trials, females preferentially took shelter in refuges scent marked by small males. Our results suggest that, like males in this species, females are able to use male scent marks to assess the body size of resident territorial males, but do not show a preference for territories occupied by large males.
Abstract: At the heart of many debates on communication is the concept of information. There is an intuitive sense in which communication implies the transfer of some kind of information, probably the reason why information is an essential ingredient in most definitions of communication. However, information has also been an endless source of misunderstandings, and recent accounts have proposed that information should be dropped from a formal definition of communication. In this article, we re-evaluate the merits and the internal logic of information-based vs. information-free approaches and conclude that information-free approaches are conceptually incomplete and operationally hindered. Instead, we propose a functional notion of information that follows logically from previous adaptationist accounts. The ensuing definition of communication provides a wider, more inclusive theoretical scope that reflects more accurately the evolutionary scenario shaping animal signals. Additionally, it is a definition better equipped to deal with the extraordinary diversity of animal signals, facilitates the distinction of honest and deceptive signals at a proximate level and accommodates a number of conceptual and practical issues (e.g. redundancy, alerting components) that are lost when we fail to acknowledge the informative content of animal signals.
Abstract: Numerosity discrimination, the ability to distinguish between sets with more and less items, is recognised as the foundation for higher numerical abilities. Understanding numerosity discrimination from a comparative perspective is hence pivotal in tracing the evolution of numerical representation systems. However, numerosity discrimination has been well studied only in vertebrates, where two innate systems of number representation have been described: an 'analog magnitude system' used to discriminate among numerosities by representing them as cardinal magnitudes and a 'parallel individualisation system' that allows precise discrimination among small arrays of items (< or =4) by representing objects individually. We investigated the existence of quantity discrimination in an insect species (Tenebrio molitor) by using a spontaneous two-choice procedure in which males were exposed to substrates bearing odours from different numbers of females (< or =4) in increasing numerosity ratios (1:4, 1:3 and 1:2). We show that males can discriminate sources of odours reflecting 1 versus 4 and 1 versus 3 females, but not 2 versus 4 or 1 versus 2, indicating that T. molitor males exhibit a marked preference for sources reflecting more female donors only when numerosity ratios are below 1:2. We discuss the functional significance of this finding and whether our pattern of results could be best explained by summation of a non-numerical continuous variable or by the existence of a numerosity discrimination mechanism with an operational signature ratio of 1:2.
Abstract: Males of many species adjust their reproductive behaviour according to the perceived risk of sperm competition. Although this phenomenon is widespread in insects and other animals, the mechanisms that allow mates to assess sperm competition levels remain largely unexplored. In this study, we analysed the mating behaviour of pairs of Tenebrio molitor beetles under three odour treatments representing increasing levels of sperm competition risk (SCR) and sperm competition intensity (SCI). Copula duration and male and female post-copulatory behaviour varied significantly with odour treatment. Both copula duration and post-copulatory associations (PCAs) increased significantly in odour treatments reflecting high male density. To our knowledge, this is the first study to report that insects may assess the actual density of potential competitors at the time of mating, a cue to SCR and SCI, on the basis of chemical cues. In T. molitor, males inhibit sperm release from the spermatophore of a rival male when remating takes place at short intervals. We show that, when sperm competition levels are high, PCAs increase female remating interval just above that necessary to prevent spermatophore inhibition by rival males. This finding strongly suggests that strategic male behaviour plays a 'spermatophore guarding' role in this species. Although common in insects with external spermatophore transfer, spermatophore guarding is not expected in species with rapid ejaculate transfer and internal spermatophore delivery. Our results reveal that spermatophore guarding may evolve, even under these circumstances, as an evolutionary response to short-term spermatophore inhibition or displacement mechanisms.
