Janne-Tuomas Seppänen

SOCIAL INFORMATION USE IS A PROCESS ACROSS TIME, SPACE, AND ECOLOGY, REACHING HETEROSPECIFICS

Decision making can be facilitated by observing other individuals faced with the same or similar problem, and recent research suggests that this social information use is a widespread phenomenon. Implications of this are diverse and profound: for example, social information use may trigger cultural evolution, affect distribution and dispersal of populations, and involve intriguing cognitive traits. We emphasize here that social information use is a process consisting of the scenes of (1) event, (2) observation, (3) decision, and (4) consequence, where the initial event is a scene in such a process of another individual. This helps to construct a sound conceptual framework for measuring and studying social information use. Importantly, the potential value of social information is affected by the distance in time, space, and ecology between the initial observation and eventual consequence of a decision. Because negative interactions between individuals (such as direct and apparent competition) also depend on the distance between individuals along these dimensions, the potential value of information and the negative interactions may form a trade-off situation. Optimal solutions to this trade-off can result in adaptively extended social information use, where using information gathered some time ago, some distance away, and from ecologically different individuals is preferred. Conceivably, using information gathered from a heterospecific individual might often be optimal. Many recent studies demonstrate that social information use does occur between species, and the first review of published cases is provided here. Such interaction between species, especially in habitat selection, has important consequences for community ecology and conservation. Adaptively extended social information use may also be an important evolutionary force in guild formation. Complex coevolutionary patterns may result depending on the effect of information use on the provider of information.

Positive fitness consequences of interspecific interaction with a potential competitor

The coexistence of species sharing mutual resources is usually thought to be limited by negative processes such as interspecific competition. This is because an overlap in resource use leads to negative fitness consequences, and traits favouring avoidance of potential competitors, for example in habitat selection, are therefore selected for. However, species interactions are acknowledged to vary from negative (competition) to mutualism, although empirical evidence for positive interspecific interactions from natural communities of other than plants and sessile animals is scarce. Here, we experimentally examined the habitat selection and its fitness consequences of a migrant bird, the pied flycatcher (Ficedula hypoleuca), in relation to the presence of competitively superior birds, resident titmice (Parus spp.). Experiments were conducted on two spatial scales: landscape and nest–site scale. We demonstrate that pied flycatchers were attracted to and accrued fitness benefits from the presence of titmice. Flycatchers breeding in tight association with titmice initiated breeding earlier, had larger broods and heavier young than solitarily breeding flycatchers. This paradoxical result indicates that species interactions may switch from negative to positive and that the coexistence of species is not always restricted by negative costs caused by other species.

Interspecific Social Learning: Novel Preference Can Be Acquired from a Competing Species

Nongenetic transmission of behavioral traits via social learning allows local traditions in humans, and, controversially, in other animals [1-4]. Social learning is usually studied as an intraspecific phenomenon (but see [5-7]). However, other species with some overlap in ecology can be more than merely potential competitors: prior settlement and longer residence can render them preferable sources of information [8]. Socially induced acquisition of choices or preferences capitalizes upon the knowledge of presumably better-informed individuals [9] and should be adaptive under many natural circumstances [10, 11]. Here we show with a field experiment that females of two migrant flycatcher species can acquire a novel, arbitrary preference of competing resident tits for a symbol attached to the nest sites. The experiment demonstrates that such blind acquisition of heterospecific traits can occur in the wild. Even though genetic variation for habitat preferences exists in many taxa [12] and overlap between bird species likely induces costs [13], this result shows that interspecific social learning can cause increased overlap in nest-site preferences. Conventional, negative species interactions push ecological niches of species apart, but the use of competing species as a source of information counters that force and may lead to convergence.

New behavioural trait adopted or rejected by observing heterospecific tutor fitness

Animals can acquire behaviours from others, including heterospecifics, but should be discriminating in when and whom to copy. Successful individuals should be preferred as tutors, while adopting traits of poorly performing individuals should be actively avoided. Thus far it is unknown if such adaptive strategies are involved when individuals copy other species. Furthermore, rejection of traits based on tutor characteristics (negative bias) has not been shown in any non-human animal. Here we test whether a choice between two new, neutral behavioural alternatives—breeding-sites with alternative geometric symbols—is affected by observing the choice and fitness of a heterospecific tutor. A field experiment replicated in four different areas shows that the proportion of pied flycatcher females matching the choice of the tit tutor consistently increased with increasing number of offspring in the tit nest, to the extent of nearly complete prevalence in one of the areas when tit fitness was highest. Notably, all four replicates demonstrate rejection of the behaviour of lowest-fitness tutors. The results demonstrate both acquisition and avoidance of heterospecific behavioural traits, based on the perceived (lack of) tutor fitness. This has potential implications for understanding the origin, diversity and local adaptations of behavioural traits, and niche overlap/partitioning and species co-occurrence.

Observed Fitness May Affect Niche Overlap in Competing Species via Selective Social Information Use

Social information transmission is important because it enables horizontal spread of behaviors, not only between conspecifics but also between individuals of different species. Because interspecific social information use is expected to take place among species with similar resource needs, it may have major consequences for the emergence of local adaptations, resource sharing, and community organization. Social information use is expected to be selective, but the conditions promoting it in an interspecific context are not well known. Here, we experimentally test whether pied flycatchers (Ficedula hypoleuca) use the clutch size of great tits (Parus major) in determining the quality of the observed individual and use it as a basis of decision making. We show that pied flycatchers copied or rejected a novel nest site feature preference of great tits experimentally manipulated to exhibit high or low fitness (clutch size), respectively. Our results demonstrate that the social transmission of behaviors across species can be highly selective in response to observed fitness, plausibly making the phenomenon adaptive. In contrast with the current theory of species coexistence, overlap between realized niches of species could dynamically increase or decrease depending on the observed success of surrounding individuals.

The past and the present in decision-making: the use of conspecific and heterospecific cues in nest site selection

Nest site selection significantly affects fitness, so adaptations for assessment of the qualities of available sites are expected. The assessment may be based on personal or social information, the latter referring to the observed location and performance of both conspecific and heterospecific individuals. Contrary to large-scale breeding habitat selection, small-scale nest site selection within habitat patches is insufficiently understood. We analyzed nest site selection in the migratory Collared Flycatcher Ficedula albicollis in relation to present and past cues provided by conspecifics and by resident tits within habitat patches by using long-term data. Collared Flycatchers preferred nest boxes that were occupied by conspecifics in the previous year. This preference was strongest in breeding pairs where both individuals bred in the same forest patch in the previous year. The results also suggest preference for nest boxes close to boxes where conspecifics had a high breeding success in the previous year, and for nest boxes which are presently surrounded by a high number of breeding Great Tits Parus major. The results indicate social information use in nest site selection at a small spatial scale, where Collared Flycatchers use conspecific cues with a time lag of one year and heterospecific cues instantly.

Observed heterospecific clutch size can affect offspring investment decisions

Optimal investment in offspring is important in maximizing lifetime reproductive success. Yet, very little is known how animals gather and integrate information about environmental factors to fine tune investment. Observing the decisions and success of other individuals, particularly when those individuals initiate breeding earlier, may provide a way for animals to quickly arrive at better breeding investment decisions. Here we show, with a field experiment using artificial nests appearing similar to resident tit nests with completed clutches, that a migratory bird can use the observed high and low clutch size of a resident competing bird species to increase and decrease clutch size and egg mass, accordingly. Our results demonstrate that songbirds can discriminate between high and low quantity of heterospecific eggs, and that social information can have long-term physiological consequences affecting reproductive strategies. Such behaviour may help animals to better adapt to changing environments and lead to convergent traits with competitors.

Intraspecific social information use in the selection of nest site characteristics

Animals commonly acquire information about the environment by monitoring how others interact with it. The importance of social information use probably varies among species. In particular, many migratory birds breeding in northern latitudes rely on social information provided by resident tits when making important decisions and are able to copy or reject selectively the decisions of tits exhibiting good or bad fitness correlates, respectively. However, little is known about the role of social information use among resident tits. In a field experiment we tested whether great tits, Parus major, given a choice between two novel alternative features on adjacent nest sites, copy or reject conspecific choice represented by a simulated tit nest nearby, when the nest exhibits a high or low fitness correlate (clutch size). In contrast to the results in the interspecific setting, the portrayed fitness of the simulated nest did not affect tits’ nest site choices. However, the age of males had an effect, irrespective of the fitness correlate. Pairs with an old male preferred, while pairs with a young male tended to reject, the nest site choices of the simulated previously settled conspecifics. This result is surprising, as the final nest site choice in this species is presumably made by females, whose age and other phenotypic traits did not have an effect here. Our results imply stronger influence of the male great tits on nest site choice than previously thought. Dominance relationships and intraspecific competition are possible explanations for the age-dependent patterns in social information use.

Avoiding perceived past resource use of potential competitors affects niche dynamics in a bird community

BackgroundSocial information use is usually considered to lead to ecological convergence among involved con- or heterospecific individuals. However, recent results demonstrate that observers can also actively avoid behaving as those individuals being observed, leading to ecological divergence. This phenomenon has been little explored so far, yet it can have significant impact on resource use, realized niches and species co-existence. In particular, the time-scale and the ecological context over which such shifts can occur are unknown. We examined with a long-term (four years) field experiment whether experimentally manipulated, species-specific, nest-site feature preferences (symbols on nest boxes) are transmitted across breeding seasons and affect future nest-site preferences in a guild of three cavity-nesting birds.ResultsOf the examined species, resident great tits (Parus major) preferred the symbol that had been associated with unoccupied nest boxes in the previous year, i.e., their preference shifted towards niche space previously unused by putative competitors and conspecifics.ConclusionsOur results show that animals can remember the earlier resource use of conspecifics and other guild members and adjust own decisions accordingly one year after. Our experiment cannot reveal the ultimate mechanism(s) behind the observed behaviour but avoiding costs of intra- or interspecific competition or ectoparasite load in old nests are plausible reasons. Our findings imply that interspecific social information use can affect resource sharing and realized niches in ecological time-scale through active avoidance of observed decisions and behavior of potentially competing species.

Peer review by the Peers, for the Peers: response to Hettyey et al.

Open discussion and participation of the scientific community are vital to the evolution of Peerage of Science and, therefore, the letter in TREE by Hettyey et al. [1] is much appreciated. Here, we reply to the three conceptual concerns that they raise. Other responses and open discussion can be found on the Peerage of Science blog (http://www.peerageofscience.org).