Variation in communicative complexity has been conceptually and empirically attributed to social complexity, with animals living in more complex social environments exhibiting more signals and/or more complex signals than animals living in simpler social environments. As compelling as studies highlighting a link between social and communicative variables are, this hypothesis remains challenged by operational problems, contrasting results, and several weaknesses of the associated tests. Specifically, how to best operationalize social and communicative complexity remains debated; alternative hypotheses, such as the role of a species' ecology, morphology, or phylogenetic history, have been neglected; and the actual ways in which variation in signaling is directly affected by social factors remain largely unexplored. In this review, we address these three issues and propose an extension of the Bsocial complexity hypothesis for communicative complexity^that resolves and acknowledges the above factors. We specifically argue for integrating the inherently multimodal nature of communication into a more comprehensive framework and for acknowledging the social context of derived signals and the potential of audience effects. By doing so, we believe it will be possible to generate more accurate predictions about which specific social parameters may be responsible for selection on new or more complex signals, as well as to uncover potential adaptive functions that are not necessarily apparent from studying communication in only one modality. Significance statement Animals exhibit an astonishing diversity of communicative systems, with important variation in both the nature and the number of signals they produce. The roles of phylogenetic history, genetic drift, environmental factors, and sexual selection in shaping interspecific variation in communicative systems have long been acknowledged, whereas social complexity has only more recently emerged as a potential fundamental factor determining signal diversification. While a number of comparative studies support the key prediction of this hypothesis, i.e., that individuals living in more complex social environments exhibit more signals and/or more complex signals, we discuss several gaps in the current state of the art concerning this hypothesis and point out what we believe are neglected perspectives. By emphasizing the importance of the multimodal nature of communicative systems and the social context in which signals are exchanged, we hope to stimulate the development of new tests and specific questions based on this expanded framework.

The origin and evolution of manual grasping remain poorly understood. The ability to cling requires important grasping abilities and is essential to survive in species where the young are carried in the fur. A previous study has suggested that this behaviour could be a pre-adaptation for the evolution of fine manipulative skills. In this study we tested the co-evolution between infant carrying in the fur and manual grasping abilities in the context of food manipulation. As strepsirrhines vary in the way infants are carried (mouth vs. fur), they are an excellent model to test this hypothesis. Data on food manipulation behaviour were collected for 21 species of strepsirrhines. Our results show that fur-carrying species exhibited significantly more frequent manual grasping of food items. This study clearly illustrates the potential novel insights that a behaviour (infant carrying) that has previously been largely ignored in the discussion of the evolution of primate manipulation can bring.

Although hand grasping is ubiquitous in primate species, its origins remain uncertain. This is in part because uncertainty about hand skills and grasping strategies persists in strepsirrhines, a monophyletic group of primates located near the base of the primate tree. In this study, we report and discuss our observations of the different grasping strategies adopted by 85 captive individuals belonging to 22 species of strepsirrhines during the grasping of food items of different sizes and consistencies. Our results indicate that although strepsirrhines do not present variability in their hand-grip types (sole whole-hand power grip), they are able to adjust their grasping strategy depending on the properties of the food. Notably, they use the mouth when more precision is needed (i.e. to grasp small items). Moreover, grasping strategies adopted for big items differ depending on food consistency, revealing a new and potentially essential factor to consider in future research on grasping abilities. We believe that by looking across this important set of species in unconstrained standardized conditions, this study provides valuable insight for further comparative research on the potential selective pressures involved in the evolution of hand grasping.