Mixed-species groups in bats: non-random roost associations and roost selection in neotropical understory bats

Background Mixed-species groups in animals have been shown to confer antipredator, foraging and other benefits to their members that may provide selective advantages. In most cases, however, it is unclear whether functional benefits are a principal driver of heterospecific groups, or whether groups simply result from simultaneous exploitation of common resources. Mixed-species groups that form independently of environmental conditions may, however, evidence direct benefits of species associations. Bats are among the most gregarious mammals, with sometimes thousands of individuals of various species roosting communally. Despite numerous potential functional benefits of such mixed-species roosting groups, interspecific attraction has never been shown. To explore alternative explanations for mixed-species roosting, we studied roost selection in a speciose neotropical understory bat community in lowland rainforest in Costa Rica. Long term roost data were recorded over 10 years in a total of 133 roosts comprising both natural roosts and structurally uniform artificial roosts. We modelled bat roost occupancy and abundance in each roost type and in forest and pasture habitats to quantify the effects of roost- and environmental variability. Results We found that bat species presence in natural roosts is predictable from habitat and structural roost parameters, but that the presence and abundance of other bat species further modifies roost choice. One third of the 12 study species were found to actively associate with selected other bat species in roosts (e.g. Glossophaga commissarisi with Carollia sowelli). Other species did not engage in communal roosting, which in some cases indicates a role for negative interspecific interactions, such as roost competition. Conclusions Mixed-species roosting may provide thermoregulatory benefits, reduce intraspecific competition and promote interspecific information transfer, and hence some heterospecific associations may be selected for in bats. Overall, our study contributes to an improved understanding of the array of factors that shape diverse tropical bat communities and drive the dynamics of heterospecific grouping in mammals more generally.

Dynamics of Cetacean Mixed-Species Groups: A Review and Conceptual Framework for Assessing Their Functional Significance

Numerous species of cetaceans have been recorded in mixed-species groups (MSGs). By forming groups with individuals of different species, cetaceans may reduce predation risk, improve foraging, and gain social benefits. Most accounts of cetacean MSGs, however, are descriptive and little is known about their functions. Furthermore, research has been hindered by inconsistent use of terminology and the lack of a conceptual framework to guide investigations. We reviewed the cetacean literature to compare how MSGs have been termed and defined, to assess their characteristics, to evaluate what is known about their potential functions, and to provide directions for future study. In total, we reviewed 203 studies reporting observations of cetacean MSGs. These MSGs involved 54 different species, predominantly delphinids, that formed 216 different species pairs with varied morphologies and levels of relatedness. Cetacean MSGs occurred across the globe, from tropical to cold temperate seas, from shallow coastal waters to the open ocean, and varied in characteristics such as group size and frequency of occurrence. Only 27 of the reviewed studies proposed and discussed the potential functions of cetacean MSGs, suggesting reduced predation risk (5 species pairs), improved foraging (17 species pairs), and social benefits (12 species pairs) as the main drivers. In most cases, however, the factors that drive the formation of cetacean MSGs remain unknown. Amongst the reviewed studies, MSGs were referred to by various terms, often with no explicit definitions. To reduce this inconsistency, we recommend that future studies use only the term mixed-species group which we define as individuals of two or more species found in close spatial proximity due to mutual or unreciprocated attraction derived from evolutionary grouping benefits. There were also few structured investigations to confirm MSG occurrence and to analyse their potential causes and consequences. To facilitate the study of cetacean MSGs, we developed a conceptual framework that establishes diverse approaches to, firstly, distinguish MSGs from chance encounters and aggregations and to, secondly, investigate their potential functions. This is necessary if we are to advance this field of study and improve our understanding of the role that MSGs play in species and community ecology.

Behavioural plasticity in a native species may be related to foraging resilience in the presence of an aggressive invader

Competition between invasive and native species can result in the exploitation of resources by the invader, reducing foraging rates of natives. However, it is increasingly recognized that multiple factors can enhance the resilience of native species competing for limiting resources with invaders. Although extensively studied in terrestrial species, little research has focused on behavioural plasticity in aquatic ecosystems and how this influences native species resilience. Here, we examined the role of behavioural plasticity in interactions between a native Australian fish, Pseudomugil signifer, and a widespread invasive fish, Gambusia holbrooki . To determine whether P. signifer displays behavioural plasticity that may mitigate competition with G. holbrooki , we first quantified social behaviours (aggression, submission and affiliation) and shoal cohesion for each species in single- and mixed-species groups. Second, we compared the feeding rates of both species in these groups to ascertain if any modulation of social behaviours and cohesion related to foraging success. We found that aggressive and submissive behaviours of G. holbrooki and P. signifer showed plasticity in the presence of heterospecifics, but social affiliation, shoaling and, most importantly, foraging, remained inflexible. This variation in the degree of plasticity highlights the complexity of the behavioural response of a native species and suggests that both behavioural modulation and consistency may be related to sustaining foraging efficiency in the presence of an invader.

Using social network analysis of mixed-species groups in African savannah herbivores to assess how community structure responds to environmental change

The dynamics of wildlife populations often depend heavily on interspecific interactions and understanding the underlying principles can be an important step in designing conservation strategies. Behavioural ecological studies can here provide useful insights into the structure and function of communities and their likely response to environmental changes. In this study of the Masai Mara herbivore community, we use a social network approach to investigate social affinities between species and how these change over the year in response to seasonal changes in ecological conditions. We find that even though social networks were correlated across different ecological conditions, for half the species dyads in the community, the strength of social affinities responded to changes in rainfall and/or the presence of migratory wildebeest. Several species consequentially adopted more or less central positions in the network depending on the ecological conditions. The findings point out interspecific social links that are likely to be attenuated or strengthened as a consequence of human-induced environmental changes and therefore call for particular attention from conservation managers. The eco-evolutionary ramifications of the perturbations of social affinities still require further study. This article is part of the theme issue ‘Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation’.

Coordinated hunting behaviors of mixed-species groups of piscivores and associated species at Isla del Coco National Park (Eastern Tropical Pacific)

ABSTRACT Studies of mixed-species groups of animals can reveal emergent complexities of collective behaviors. In this study we collected data on mixed-species hunting groups composed primarily of piscivorous fishes (species composition, abundance, behavioral interactions) and used both multivariate and network analyses to quantify pair-wise and guild level behavioral relationships. Our results indicate that such collective behaviors exhibit consistent patterns of associations (33 species with 282 pair-wise links within the observed network) with 10 dominant species accounting for 60% of pair-wise interactions. Species richness within groups varied (mean = 2.4, range 2-6 species) as did group size (mean = 8.1 individuals, range 2-80). Mixed-species groups, in general, were composed of species representing morphologically diverse forms that appeared to enhance access to shelter sites and implement diverse strategies for prey capture. It is noteworthy that the composition of groups did not reflect the relative abundances of their component species within the overall community of fishes, suggesting that group membership was an elective choice. The identification of these patterns, assuming they are persistent features of these communities, can be used as a foundation for studies to assess dynamics of mixed-species relationships, rates of predator success based on group membership, demographic consequences, and responses to variations in habitat attributes and associated prey resources. Such information could be used to interpret the nature of multispecies interactions within predator communities and potentially aid in conservation and management.

Species-specific strategies increase unpredictability of escape flight in eared moths

ABSTRACTMany prey animals form mixed-species groups. Mixed-species groups provide various benefits ranging increased food intake to increased chance of predator detection. The escape-tactic diversity hypothesis predicts another benefit. It postulates that the overall unpredictability of evasive movement is increased if multiple species with different evasive tactics mix, resulting in enhanced predator protection for the whole group. Echolocating bats and eared moths are a textbook example of predator-prey interactions. Moths exhibit evasive flight with diverse tactics; however, the variability of their evasive flight within and between species has never been systematically quantified. We therefore recorded flight strength of eight moth species in response to the same level of simulated bat predation. We show species-specific and size-independent differences in both overall flight strength and in change of flight strength over time, confirming the escape-tactic diversity hypothesis for eared moths. Additionally, we show strong inter-individual differences in evasive flight within some species. This diversity in escape tactic between eared moths increases the overall unpredictability of their evasive flight in mixed-species groups, likely providing better protection against predatory bats for the single individual.

Cohesion, order and information flow in the collective motion of mixed-species shoals

Despite the frequency with which mixed-species groups are observed in nature, studies of collective behaviour typically focus on single-species groups. Here, we quantify and compare the patterns of interactions between three fish species, threespine sticklebacks ( Gasterosteus aculeatus ), ninespine sticklebacks ( Pungitius pungitius ) and roach ( Rutilus rutilus ) in both single- and mixed-species shoals in the laboratory. Pilot data confirmed that the three species form both single- and mixed-species shoals in the wild. In our laboratory study, we found that single-species groups were more polarized than mixed-species groups, while single-species groups of threespine sticklebacks and roach were more cohesive than mixed shoals of these species. Furthermore, while there was no difference between the inter-individual distances between threespine and ninespine sticklebacks within mixed-species groups, there was some evidence of segregation by species in mixed groups of threespine sticklebacks and roach. There were differences between treatments in mean pairwise transfer entropy, and in particular we identify species-differences in information use within the mixed-species groups, and, similarly, differences in responses to conspecifics and heterospecifics in mixed-species groups. We speculate that differences in the patterns of interactions between species in mixed-species groups may determine patterns of fission and fusion in such groups.