Ecological and evolutionary consequences of selective interspecific information use.

Recent work has shown that animals frequently use social information from individuals of their own species as well as from other species; however, the ecological and evolutionary consequences of this social information use remain poorly understood. Additionally, information users may be selective in their social information use, deciding from whom and how to use information, but this has been overlooked in an interspecific context. In particular, the intentional decision to reject a behaviour observed via social information has received less attention, although recent work has indicated its presence in various taxa. Based on existing literature, we explore in which circumstances selective interspecific information use may lead to different ecological and coevolutionary outcomes between two species, such as explaining observed co-occurrences of putative competitors. The initial ecological differences and the balance between the costs of competition and the benefits of social information use potentially determine whether selection may lead to trait divergence, convergence or coevolutionary arms race between two species. We propose that selective social information use, including adoption and rejection of behaviours, may have far-reaching fitness consequences, potentially leading to community-level eco-evolutionary outcomes. We argue that these consequences of selective interspecific information use may be much more widespread than has thus far been considered.

Foraging Bumblebees Selectively Attend to Other Types of Bees Based on Their Reward-Predictive Value.

Using social information can be an efficient strategy for learning in a new environment while reducing the risks associated with trial-and-error learning. Whereas social information from conspecifics has long been assumed to be preferentially attended by animals, heterospecifics can also provide relevant information. Because different species may vary in their informative value, using heterospecific social information indiscriminately can be ineffective and even detrimental. Here, we evaluated how selective use of social information might arise at a proximate level in bumblebees (Bombus terrestris) as a result of experience with demonstrators differing in their visual appearance and in their informative value as reward predictors. Bumblebees were first trained to discriminate rewarding from unrewarding flowers based on which type of "heterospecific" (one of two differently painted model bees) was next to each flower. Subsequently, these bumblebees were exposed to a novel foraging context with two live painted bees. In this novel context, observer bumblebees showed significantly more social information-seeking behavior towards the type of bees that had predicted reward during training. Bumblebees were not attracted by paint-marked small wooden balls (moved via magnets) or paint-marked non-pollinating heterospecifics (woodlice; Porcellio laevis) in the novel context, indicating that bees did not simply respond to conditioned color cues nor to irrelevant social cues, but rather had a "search image" of what previously constituted a valuable, versus invaluable, information provider. The behavior of our bumblebees suggests that their use of social information is governed by learning, is selective, and extends beyond conspecifics.

Interspecific information on predation risk affects nest site choice in a passerine bird.

BACKGROUND: Breeding site choice constitutes an important part of the species niche. Nest predation affects breeding site choice, and has been suggested to drive niche segregation and local coexistence of species. Interspecific social information use may, in turn, result in copying or rejection of heterospecific niche characteristics and thus affect realized niche overlap between species. We tested experimentally whether a migratory bird, the pied flycatcher Ficedula hypoleuca, collects information about nest predation risk from indirect cues of predators visiting nests of heterospecific birds. Furthermore, we investigated whether the migratory birds can associate such information with a specific nest site characteristic and generalize the information to their own nest site choice. RESULTS: Our results demonstrate that flycatchers can use the fate of heterospecific nesting attempts in their own nest site choice, but do so selectively. Young flycatcher females, when making the decision quickly, associated the fate of an artificial nest with nest-site characteristics and avoided the characteristic associated with higher nest predation risk. CONCLUSIONS: Copying nest site choices of successful heterospecifics, and avoiding choices which led to failed attempts, may amplify or counter effects of nest predation on niche overlap, with important consequences for between-species niche divergence-convergence dynamics, species coexistence and predator-prey interactions.

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

BACKGROUND: Social 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. RESULTS: Of 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. CONCLUSIONS: Our 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.

Active hiding of social information from information-parasites.

BACKGROUND: Coevolution between pairs of different kind of entities, such as providers and users of information, involves reciprocal selection pressures between them as a consequence of their ecological interaction. Pied flycatchers (Ficedula hypoleuca) have been shown to derive fitness benefits (larger clutches) when nesting in proximity to great tits (Parus major), presumably because they this way discover and obtain information about nesting sites. Tits suffer from the resulting association (smaller clutches). An arms race between the tits (information host) and the flycatchers (information parasite) could thus result. Great tits often cover eggs with nesting material before, but not during incubation. We hypothesized that one function of egg-covering could be a counter-adaptation to reduce information parasitism by pied flycatchers. We predicted that tits should bring more new hair to cover their exposed eggs when a pied flycatcher is present near to tit nest than when a neutral (non-competing) species is present. We conducted decoy and playback experiment in Oulu and Turku, Finland. First, we removed and collected all the hair covering the tit eggs. Then, we measured how the perceived presence of flycatcher or waxwing (Bombycilla garrulus) affects tits' egg-covering by collecting and weighing the hair brought on the eggs and photographing the nest 24 h after the playback. RESULTS: Tits brought more hair into the nest and covered the eggs more carefully after flycatcher treatment, compared to waxwing treatment. We also found that the tits in Oulu (over 600 km to north from Turku) had more hair on the top of their eggs in general. CONCLUSIONS: Together, these results suggest that the counter-adaptation function of egg-covering against information parasites may be an extension of original function to protect eggs from low temperatures.

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.

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.

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.

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.

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.