Robert F. Lachlan

Who follows whom? Shoaling preferences and social learning of foraging information in guppies.

Preferences of fish for different types of shoals may influence the transmission of novel information through them. We investigated the factors influencing the preferences of guppies, Poecilia reticulata, for different shoals in order to shed some light on how information transmission occurs. Adult subjects were given a choice between swimming with two diverging shoals of conspecifics that differed with respect to key characteristics. In six choice experiments, subjects discriminated between shoal partners on the basis of: (1) shoal size, subjects preferring a shoal of 10 to a single fish; (2) size of shoaling fish, small fish preferring small conspecifics rather than an equal number of large fish, while large fish showed no preference; (3) local foraging experience of shoaling fish, shoals containing fish that had previously been repeatedly fed in the experimental tank being preferred to shoals with no such experience; and (4) familiarity of shoaling fish, guppies preferring familiar rather than unfamiliar conspecifics. No discrimination on the basis of colour or hunger was observed. In addition, following a shoal to a food site on just three trials allowed guppies to learn a route, or food site, preference. Guppies were considerably more likely to learn to adopt the behaviour shown by members of a shoal of several demonstrators than an alternative behaviour shown by a single conspecific demonstrator. The relationship between preferences for different shoals and the social transmission of information is discussed in the light of these findings. The results suggest that shoaling preferences may strongly influence the social transmission of novel foraging information or feeding preferences through fish populations, and imply that learned infor-mation may diffuse through fish populations in a nonrandom, or directed, manner. Copyright 1998 The Association for the Study of Animal Behaviour.

SONG LEARNING ACCELERATES ALLOPATRIC SPECIATION

Abstract The songs of many birds are unusual in that they serve a role in identifying conspecific mates, yet they are also culturally transmitted. Noting the apparently high rate of diversity in one avian taxon, the songbirds, in which song learning appears ubiquitous, it has often been speculated that cultural transmission may increase the rate of speciation. Here we examine the possibility that song learning affects the rate of allopatric speciation. We construct a population‐genetic model of allopatric divergence that explores the evolution of genes that underlie learning preferences (predispositions to learn some songs over others). We compare this with a model in which mating signals are inherited only genetically. Models are constructed for the cases where songs and preferences are affected by the same or different loci, and we analyze them using analytical local stability analysis combined with simulations of drift and directional sexual selection. Under nearly all conditions examined, song divergence occurs more readily in the learning model than in the nonlearning model. This is a result of reduced frequency‐dependent selection in the learning models. Cultural evolution causes males with unusual genotypes to tend to learn from the majority of males around them, and thus develop songs compatible with the majority of the females in the population. Unusual genotypes can therefore be masked by learning. Over a wide range of conditions, learning therefore reduces the waiting time for speciation to occur and can be predicted to accelerate the rate of speciation.

FEMALE MATE‐CHOICE BEHAVIOR AND SYMPATRIC SPECIATION

Abstract Many models have investigated how the process of speciation may occur in sympatry. In these models, individuals are either asexual or mate choice is determined by very simple rules. Females, for example, may be assumed either to compare their phenotype to that of a potential mate, preferring to mate with similar males (phenotype matching), or to possess preference genes that determine which male phenotype they prefer. These rules often do not reflect the mate-choice rules found in empirical studies. In this paper, we compare these two modes of female choice with various types of sexual imprinting. We examine the efficacy of different mate-choice behavior in causing divergence in male traits under simple deterministic one-locus population genetic models as well as under polygenic, individual-based simulations based on the models of Dieckmann and Doebeli (1999). We find that the inheritance mechanism of mate choice can have a large effect on the ease of sympatric speciation. When females imprint on their mothers, the result of the model is similar to phenotype matching, where speciation can occur fairly easily. When females imprint on their fathers or imprint obliquely, speciation becomes considerably less likely. Finally, when females rely on preference genes, male trait evolution occurs easily, but the correlation between trait and preference can be weak, and interpreting these results as speciation may be suspect.

The maintenance of vocal learning by gene-culture interaction: the cultural trap hypothesis

Vocal learning has evolved in several groups of animals, yet the reasons for its origins and maintenance are controversial, with none of the theories put forward appearing to apply over a broad range of species. The theory of gene–culture coevolution is applied to this problem taking the specific case of the maintenance of song learning in birds. The interaction between genes underlying the filter for recognizing and learning conspecific song and the culturally transmitted songs themselves sets up an evolutionary force that may maintain vocal learning. We evaluate this hypothesis using a spatial simulation model. Our results suggest that selection that would maintain song learning exists over a wide range of conditions. Song learning may persist due to an evolutionary trap even though the average fitness in a population of learners may be lower than in a population of non–learners.

Social learning directs feeding preferences in the zebra finch, Taeniopygia guttata

Abstract We tested sexually mature zebra finches to see whether social learning influenced their feeding preferences, in particular whether they followed the preference of a male or a female demonstrator, of a red-ringed or a green-ringed male, and of a familiar or an unfamiliar male. Each observer was exposed to two demonstrators feeding at different-coloured hoppers, and then tested with a choice of hoppers to see which of the two colours they preferred. Males showed no preference between male and female demonstrators when choosing from which colour of food hopper to feed, but females preferred to feed from the hopper colour the male demonstrator had used. Both males and females exposed to male demonstrators wearing red or green leg rings fed preferentially from the same colour hopper as the red-ringed demonstrators had used. Finally, male birds exposed to familiar and unfamiliar demonstrators, preferred the food hopper from which the familiar demonstrator had fed. We interpret the results as indicating differences between the demonstrators in the amount of attention they attracted from observers. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

The evolution of conformity-enforcing behaviour in cultural communication systems

Conformity is an important aspect of many communication systems in which signals are culturally transmitted. We suggest that one reason for it to evolve is if nonconforming individuals are discriminated against, and we therefore investigated how this might arise. We designed a spatial simulation, in which individuals occupied a territory on a lattice, and played a territorial game with their neighbours in which one individual could challenge another and, if successful, obtain some of the latters resources. We then examined the relative success of pairs of strategies. We found that aggression targeted towards nonconformers (conformity-enforcing behaviour) was more successful than randomly targeted aggression in both a simple model and a more complicated one in which the conditions were based on the specific case of song sharing between songbirds. The reason for this result is that individuals could use patterns of vocal sharing to gang up on neighbours with locally uncommon song types, increasing their own chances of success. In both models, a critical parameter was how the number of contests an individual was previously involved in altered the chance of winning: conformity-enforcing behaviour was the most successful strategy only if the more contests an individual had been involved in, the less likely it was to win. Our results may explain the origin of conformity-enforcing behaviour and discrimination in nonhuman animals and may have relevance to similar behaviour in humans.

Evolution of cultural communication systems: the coevolution of cultural signals and genes encoding learning preferences.

In several communication systems that rely on social learning, such as bird song, and possibly human language, the range of signals that can be learned is limited by perceptual biases – predispositions – that are presumably based on genes. In this paper, we examine the coevolution of such genes with the culturally transmitted communication traits themselves, using deterministic population genetic models. We argue that examining how restrictive genetic predispositions are is a useful way of examining the evolutionary origin and maintenance of learning. Under neutral cultural evolution, where no cultural trait has any inherent advantage over another, there is selection in favour of less restrictive genes (genes that allow a wider range of signals to recognized). In contrast, cultural conformity (where the most common cultural trait is favoured) leads to selection in favour of more restrictive genes.

Are there species-universal categories in bird song phonology and syntax? A comparative study of chaffinches (Fringilla coelebs), zebra finches (Taenopygia guttata), and swamp sparrows (Melospiza georgiana).

Previous studies of learned bird song have suggested the existence of species-universal patterns in song organization: clear clusters in produced songs that do not vary within a species. Here the authors combine a computational method of comparing songs with statistical methods of assessing cluster structure to investigate this issue in a more quantitative manner. The authors first analyze song phonology and then examine song syntax at a population level in 3 species with very different song structure: chaffinches (Fringilla coelebs), zebra finches (Taenopygia guttata), and swamp sparrows (Melospiza georgiana). The authors used a dynamic time-warping algorithm to compare song elements, which closely matched the judgments of human observers. Clustering tendency and validation statistics showed that broad phonological categories existed in all 3 species, but these categories explained no more than half of the overall phonological variation. The authors developed a novel statistic to assess syntactical structure, which indicated that element transitions were not randomly distributed. In the clearest case, in chaffinches, this could be explained by syllables being linked to certain positions within the song. These results demonstrate measures of song organization that can be applied across species, enhancing the potential of comparative studies.

Speed-accuracy trade-offs and individually consistent decision making by individuals and dyads of zebrafish in a colour discrimination task

M.Y.W. was supported by the Overseas Research Student Awards Scheme and the Ministry of Education and National Science Council Taiwan Studying Abroad Scholarship (grant number 1003116013).

How Reliable Is Song Learning Accuracy as a Signal of Male Early Condition

That many species of songbirds learn their songs imitatively is well established, but it is less clear why they do so. A component of the developmental-stress hypothesis posits that young males in good condition learn songs more accurately than males in poor condition and that females use learning accuracy as an honest signal of male developmental history. An unresolved problem is how females reliably assess learning accuracy when they are not certain of the identity of the male’s tutor and thus the specific model from which a song was copied. We therefore investigated whether song learning accuracy assessment (SLAA) can be reliable, using evolutionary simulation models of song learning. We found that SLAA is indeed less reliable than assessment in which male signals are compared to an unlearned standard, as a result of three types of errors in matching songs to their models. In the simplest models, SLAA was particularly unreliable, but when the model is made more realistic by including features such as geographically constrained learning, repertoire complexity, and, in particular, song categorization, the reliability of SLAA increased. Our results demonstrate a range of conditions under which the assessment of song learning accuracy might be reasonably reliable and therefore likely to evolve.