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.

Group living is a critical component of the ecology of social animals such as delphinids. In many studies on these animals, groups represent sampling units that form the basis of the collection and analysis of data on their abundance, behavior, and social structure. Nevertheless, defining what constitutes a group has proven problematic. There is inconsistency in the terms and criteria used and many definitions lack biological justification. We conducted a literature review and an online expert survey to assess various terms ( group , school , party , and pod ), and their definitions as applied to delphinids to identify issues to ultimately make recommendations. Of 707 studies analyzed, 325 explicitly defined one or more terms, providing 344 definitions. Additionally, 192 definitions were obtained from the survey. Among these definitions, group was the most common term used (review: 286 definitions, 83.1%; survey: 69 definitions, 35.9%) and the most familiar to the survey respondents (73 respondents, 100.0%). In definitions of group , spatial proximity was the most used criterion (review: 200 definitions, 71.2%; survey: 53 definitions, 81.5%) followed by behavior (review: 176 definitions, 62.6%; survey: 38 definitions, 58.5%). The terms and criteria used to define delphinid groups vary considerably. Rather than proposing a single formal definition, we instead recommend that the term group and spatial proximity criteria be used to define sampling units of individuals observed in the field. Furthermore, we propose a process for formulating definitions that involves analyzing interindividual distances to determine naturally occurring patterns that are indicative of group membership. Although this process is based principally on the spatial proximity of individuals, it could also incorporate the behavior of group members by evaluating the influence of behavior on interindividual distances. Such a process produces definitions that are biologically meaningful and compatible across studies and populations, thus increasing our ability to draw strong conclusions about group living in delphinids.

Mixed-species groups are common amongst diverse taxa including fishes, birds, and mammals. Antipredator, foraging, and social benefits have been proposed as functional explanations for mixed-species group formation. Amongst delphinids, mixed-species groups are widespread, but little is known about their function. To investigate the potential benefits of delphinid mixed-species groups, we compared the number of individuals, the age composition, and the behaviour of single- and mixed-species sightings of Australian humpback (Sousa sahulensis) and Indo-Pacific bottlenose dolphins (Tursiops aduncus) observed around the North West Cape, Western Australia. We found no difference in the number of individuals or the age composition of humpback dolphins present in single- and mixed-species sightings, whereas bottlenose dolphins were present in larger numbers in single-species sightings than in mixed-species sightings due to a higher number of adults. Socialising was the initial observed behavioural state of 36.1% of mixed-species sightings, compared to only 5.1% and 10.3% of humpback and bottlenose dolphin single-species sightings, respectively. Furthermore, both species travelled and foraged less frequently when in mixed-species groups. Of 93 mixed-species groups observed during a focal follow of ≥ 10 min, 32 (34.4%) involved aggressive and/or sexual behaviours typically initiated by bottlenose dolphins towards humpback dolphins while the remaining 61 (65.6%) involved only neutral and affiliative behaviours. The results of this study suggest that the observed mixed-species groups provide multiple social benefits, particularly those pertaining to socio-sexual behaviours and the development and care of young. Significance statement Numerous species of dolphin are known to form mixed-species groups. Few studies have, however, investigated the antipredator, foraging, and/or social benefits that these species may gain by grouping with other species. Investigating these benefits is key to understanding the impacts of mixed-species groups on the species and individuals involved. We evaluated the potential functions of mixed-species groups of two sympatric, inshore dolphin species — the Australian humpback dolphin and the Indo-Pacific bottlenose dolphin. We found that both species engage in socialising behaviour more frequently when in mixed-species groups and that they engage in a diverse variety of interspecific behavioural interactions. Our results not only indicate that social benefits are the key drivers of these mixed-species groups, but they also highlight the complexity of social interactions between dolphin species.

Numerous species have been reported to form mixed-species groups, however, little is known about the interplay between niche partitioning and mixed-species group formation. Furthermore, it is often unclear whether species come together by chance due to overlapping habitat preferences, by shared attraction to resources, or by attraction between them. We assessed habitat partitioning, co-occurrence patterns, and mixed-species group formation of sympatric Australian humpback (Sousa sahulensis) and Indo-Pacific bottlenose dolphins (Tursiops aduncus) around the North West Cape, Western Australia, with a joint species distribution model and temporal analyses of sighting data. Australian humpback dolphins preferred shallower and more nearshore waters than Indo-Pacific bottlenose dolphins, yet these species co-occurred more often than expected by chance given shared responses to environmental variables. Indo-Pacific bottlenose dolphins were sighted more often than Australian humpback dolphins during the afternoon, however, we did not find any temporal patterns in the occurrence of mixed-species groups. We propose that the positive association in the species’ occurrence indicates the active formation of mixed-species groups. By evaluating habitat partitioning and co-occurrence patterns, this study provides direction for future work which should proceed to investigate the benefits that these species may gain from grouping with each other.