Comparing single- and mixed-species groups in fruit flies: differences in group dynamics, but not group formation.

Mixed-species groups describe active associations among individuals of 2 or more species at the same trophic level. Mixed-species groups are important to key ecological and evolutionary processes such as competition and predation, and research that ignores the presence of other species risks ignoring a key aspect of the environment in which social behavior is expressed and selected. Despite the defining emphasis of active formation for mixed-species groups, surprisingly little is known about the mechanisms by which mixed-species groups form. Furthermore, insects have been almost completely ignored in the study of mixed-species groups, despite their taxonomic importance and relative prominence in the study of single-species groups. Here, we measured group formation processes in Drosophila melanogaster and its sister species, Drosophila simulans. Each species was studied alone, and together, and one population of D. melanogaster was also studied both alone and with another, phenotypically distinct D. melanogaster population, in a nested-factorial design. This approach differs from typical methods of studying mixed-species groups in that we could quantitatively compare group formation between single-population, mixed-population, and mixed-species treatments. Surprisingly, we found no differences between treatments in the number, size, or composition of groups that formed, suggesting that single- and mixed-species groups form through similar mechanisms of active attraction. However, we found that mixed-species groups showed elevated interspecies male-male interactions, relative to interpopulation or intergenotype interactions in single-species groups. Our findings expand the conceptual and taxonomic study of mixed-species groups while raising new questions about the mechanisms of group formation broadly.

Effect of density and species preferences on collective choices: an experimental study on maggot aggregation behaviours.

Collective decisions have been extensively studied in arthropods, but they remain poorly understood in heterospecific groups. This study was designed to (1) assess the collective behaviours of blow fly larvae (Diptera: Calliphoridae) in groups varying in density and species composition, and (2) relate them to the costs and benefits of aggregating on fresh or decomposed food. First, experiments testing conspecific groups of Lucilia sericata and Calliphora vicina larvae, two species feeding at the same time on fresh carcasses, demonstrated decreases in growth and survival on rotten beef liver compared with fresh liver. However, mixing species together reduced this adverse impact of decomposition by increasing the mass of emerged adults. Second, larval groups were observed in binary choice tests between fresh and rotten liver (i.e. optimal and sub-optimal food sources). The results showed that larvae interacted with each other and that these interactions influenced their food preferences. We observed that (1) larvae were able to collectively choose the optimal food, (2) their choice accuracy increased with larval density and (3) the presence of another species induced a reversal in larval preference towards rotten food. These results highlight the ubiquity of collective decision properties in gregarious insects. They also reveal an unexpected effect of interspecific association, suggesting the colonization of new resources through a developmental niche construction.

Interspecific shared collective decision-making in two forensically important species.

To date, the study of collective behaviour has mainly focused on intraspecific situations: the collective decision-making of mixed-species groups involving interspecific aggregation-segregation has received little attention. Here, we show that, in both conspecific and heterospecific groups, the larvae of two species (Lucilia sericata and Calliphora vomitoria, calliphorid carrion-feeding flies) were able to make a collective choice. In all groups, the choice was made within a few minutes and persisted throughout the period of the experiment. The monitoring of a focal individual within a group showed that these aggregations were governed by attractive and retentive effects of the group. Furthermore, the similarity observed between the conspecific and heterospecific groups suggested the existence of shared aggregation signals. The group size was found to have a stronger influence than the species of necrophagous larvae. These results should be viewed in relation to the well-known correlation between group size and heat generation. This study provides the first experimental examination of the dynamics of collective decision-making in mixed-species groups of invertebrates, contributing to our understanding of the cooperation-competition phenomenon in animal social groups.

Behavioural hypervolumes of spider communities predict community performance and disbandment.

Trait-based ecology argues that an understanding of the traits of interactors can enhance the predictability of ecological outcomes. We examine here whether the multidimensional behavioural-trait diversity of communities influences community performance and stability in situ We created experimental communities of web-building spiders, each with an identical species composition. Communities contained one individual of each of five different species. Prior to establishing these communities in the field, we examined three behavioural traits for each individual spider. These behavioural measures allowed us to estimate community-wide behavioural diversity, as inferred by the multidimensional behavioural volume occupied by the entire community. Communities that occupied a larger region of behavioural-trait space (i.e. where spiders differed more from each other behaviourally) gained more mass and were less likely to disband. Thus, there is a community-wide benefit to multidimensional behavioural diversity in this system that might translate to other multispecies assemblages.

Interspecific signalling between mutualists: food-thieving drongos use a cooperative sentinel call to manipulate foraging partners.

Interspecific communication is common in nature, particularly between mutualists. However, whether signals evolved for communication with other species, or are in fact conspecific signals eavesdropped upon by partners, is often unclear. Fork-tailed drongos (Dicrurus adsimilis) associate with mixed-species groups and often produce true alarms at predators, whereupon associating species flee to cover, but also false alarms to steal associating species' food (kleptoparasitism). Despite such deception, associating species respond to drongo non-alarm calls by increasing their foraging and decreasing vigilance. Yet, whether these calls represent interspecific sentinel signals remains unknown. We show that drongos produced a specific sentinel call when foraging with a common associate, the sociable weaver (Philetairus socius), but not when alone. Weavers increased their foraging and decreased vigilance when naturally associating with drongos, and in response to sentinel call playback. Further, drongos sentinel-called more often when weavers were moving, and weavers approached sentinel calls, suggesting a recruitment function. Finally, drongos sentinel-called when weavers fled following false alarms, thereby reducing disruption to weaver foraging time. Results therefore provide evidence of an 'all clear' signal that mitigates the cost of inaccurate communication. Our results suggest that drongos enhance exploitation of a foraging mutualist through coevolution of interspecific sentinel signals.

Connecting and integrating cooperation within and between species.

There has long been a fundamental divide in the study of cooperation: researchers focus either on cooperation within species, including but not limited to sociality, or else on cooperation between species, commonly termed mutualism. Here, we explore the ecologically and evolutionarily significant ways in which within- and between-species cooperation interact. We highlight two primary cross-linkages. First, cooperation of one type can change the context in which cooperation of the other type functions, and thus potentially its outcome. We delineate three possibilities: (i) within-species cooperation modulates benefits for a heterospecific partner; (ii) between-species cooperation affects the dynamics of within-species cooperation; and (iii) both processes take place interactively. The second type of cross-linkage emerges when resources or services that cooperation makes available are obtainable either from members of the same species or from different species. This brings cooperation at the two levels into direct interaction, to some extent obscuring the distinction between them. We expand on these intersections between within- and between-species cooperation in a diversity of taxa and interaction types. These interactions have the potential to weave together social networks and trophic dynamics, contributing to the structure and functioning of ecological communities in ways that are just beginning to be explored. This article is part of the theme issue 'Connected interactions: enriching food web research by spatial and social interactions'.

Do mixed-species groups of capuchin (Sapajus apella) and squirrel monkeys (Saimiri sciureus) synchronize their behaviour?

In the wild, coordinated behaviour across group members is essential for maintaining spatial coherence, with potential implications for individual fitness. Such coordination often leads to behavioural synchrony (performing the same behaviour at the same time). Tufted capuchins (Sapajus apella) and squirrel monkeys (Saimiri sciureus) are known to form mixed-species groups (MSGs), travelling and foraging together. Yet, it is unclear if it is necessary to synchronize behaviours in captivity when ecological pressures are minimal compared to the wild. We investigated the extent to which two MSGs of capuchins (N = 35) and squirrel monkeys (N = 26) synchronized their behaviour with conspecifics and heterospecifics at the Living Links to Human Evolution Research Centre, RZSS, Edinburgh Zoo, UK. Group activities were sampled by instantaneous scans of all visible individuals. Scans (n = 180) were analysed for five most frequently observed behaviours. Intraspecies synchrony was calculated using Simpson's Diversity Index, and interspecies synchrony was measured using cross-correlations. Intraspecific synchrony was significantly greater compared to randomly aggregated data, while cross-correlations indicated interspecific asynchrony. Living together did not lead to interspecific synchrony as may be expected given the coordination and behaviour described in the wild, and shared husbandry in captivity. Overall, our findings highlight differences in the behavioural structure of single- versus MSGs. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

A mixed-species shoal including fewer heterospecifics is preferred by jack mackerel Trachurus japonicus juveniles.

Mixed-species shoals of fishes were reported from various environments such as coral reefs and temperate lakes. The driving mechanism of forming mixed-species shoal can be either haphazard or a preferential choice by an individual. Here, we tested if jack mackerel Trachurus japonicus, a temperate marine pelagic fish, could choose a shoal composition using vision based on a choice experiment. Shoal preference of jack mackerel juveniles was evaluated by presenting shoals comprising one to four heterospecifics (half-lined cardinal Ostorhinchus semilineatus) out of four shoal members to the focal fish. When both conspecific and heterospecific shoals were presented to a single jack mackerel, the focal fish showed a preference for the conspecific rather than heterospecific shoal, indicating that they can choose a conspecific shoal based on the visual information. However, when a focal fish encountered both conspecific and mixed-species shoals including fewer heterospecifics (one out of four individuals), the latter was preferred. The result implies that jack mackerel can judge a preferable composition of shoal using visual information. The inclusion of fewer heterospecifics may be exploiting oddity effects, whereby minority species are exposed to relatively high vulnerability to predators, as well as resulting in reduced competition for food while maintaining predator vigilance in a shoal.

Friendship across species borders: factors that facilitate and constrain heterospecific sociality.

Our understanding of animal sociality is based almost entirely on single-species sociality. Heterospecific sociality, although documented in numerous taxa and contexts, remains at the margins of sociality research and is rarely investigated in conjunction with single-species sociality. This could be because heterospecific and single-species sociality are thought to be based on fundamentally different mechanisms. However, our literature survey shows that heterospecific sociality based on mechanisms similar to single-species sociality is reported from many taxa, contexts and for various benefits. Therefore, we propose a conceptual framework to understand conspecific versus heterospecific social partner choice. Previous attempts, which are all in the context of social information, model partner choice as a trade-off between information benefit and competition cost, along a single phenotypic distance axis. Our framework of partner choice considers both direct grouping benefits and information benefits, allows heterospecific and conspecific partners to differ in degree and qualitatively, and uses a multi-dimensional trait space analysis of costs (competition and activity matching) and benefits (relevance of partner and quality of partner). We conclude that social partner choice is best-viewed as a continuum: some social benefits are obtainable only from conspecifics, some only from dissimilar heterospecifics, while many are potentially obtainable from conspecifics and heterospecifics.This article is part of the theme issue 'Collective movement ecology'.

Voice discrimination in four primates.

One accepted function of vocalisations is to convey information about the signaller, such as its age-sex class, motivation, or relationship with the recipient. Yet, in natural habitats individuals not only interact with conspecifics but also with members of other species. This is well documented for African forest monkeys, which form semi-permanent mixed-species groups that can persist for decades. Although members of such groups interact with each other on a daily basis, both physically and vocally, it is currently unknown whether they can discriminate familiar and unfamiliar voices of heterospecific group members. We addressed this question with playbacks on monkey species known to form polyspecific associations in the wild: red-capped mangabeys, Campbell's monkeys and Guereza colobus monkeys. We tested subjects' discrimination abilities of contact calls of familiar and unfamiliar female De Brazza monkeys. When pooling all species, subjects looked more often towards the speaker when hearing contact calls of unfamiliar than familiar callers. When testing De Brazza monkeys with their own calls, we found the same effect with the longest gaze durations after hearing unfamiliar voices. This suggests that primates can discriminate, not only between familiar and unfamiliar voices of conspecifics, but also between familiar and unfamiliar voices of heterospecifics living within a close proximity.