When inter-individual differences in behavioural traits are consistent across time and/or context they are considered personality traits ( Réale and Dingemanse, 2012). It is indeed unknown whether some individual characteristics of workers, such as personality traits, would predict tool use behaviour. We asked the question of what makes a good tool user in A. Moreover, there was no significant relationship between the number of ants working at the debris dropping task and the number of debris pieces dropped, indicating that a small number of workers can perform this task very efficiently ( Lőrinczi, 2014). The number of tool users did not increase with colony size, while the number of total foragers did. subterranea as well, the proportion of workers observed to use tools was only a small fraction of the total number of foragers. rudis, it was observed that the tool use behaviour is carried out by a small subset of individuals within the group of foragers and only a small number of workers perform the debris dropping task ( Banschbach et al., 2006). In particular, we do not know whether any forager with sufficient information about the location of the food and the availability of the tools would perform tool use or whether there are specialised workers which perform tool use repeatedly. The process of tool use in ants has not been studied at the individual level. Tool selection also depends on the foraging environment and varies with food type (viscosity), distance, and availability of tools ( Lőrinczi et al., 2018). Furthermore, ants can learn to use artificial material that is novel to them and select the material with optimal soaking properties, thus showing that tool use is not behaviourally fixed in ants ( Maák et al., 2017). In a previous study ( Maák et al., 2017), we have investigated whether Aphaenogaster ants are selective in the choice of material to be used as tools and we demonstrated that ant workers prefer materials that are easy to handle and with good soaking capacity. Indeed, these ants do not drop debris in non-food substances ( Banschbach et al., 2006). This behaviour qualifies as tool use, namely tool-assisted food transport. When Aphaenogaster foragers discover a source of liquid food, such as fruit pulp or body fluids of dead insects, they collect debris (pieces of leaves, soil, sand grains), drop them into the liquid food, and then transport these soaked debris to their nest. Workers forage individually mostly on the ground level and can cover large areas in habitats with scarce food sources ( Cerdá et al., 1998). trophallaxis, Delage and Jaisson, 1969) therefore, foragers cannot use this form of food transmission among colony members. Moreover, unlike most ant species, Aphaenogaster workers do not perform mouth-to-mouth exchange of liquid food (i.e. Workers of these species are characterised by the lack of a distensible crop and by a chitinous gaster, preventing the transportation of large amounts of liquid food inside their bodies, a feature common in many other ant species ( Hölldobler and Wilson, 1990 Davidson et al., 2004). Among the best described examples is the use of debris to transport liquid food by some species of ants ( Morrill, 1972 Barber et al., 1989), in particular, several species of the genus Aphaenogaster ( Fellers and Fellers, 1976 Tanaka and Ono, 1978 McDonald, 1984 Agbogba, 1985). Smith and Bentley-Condit, 2010 reported about 50 cases of tool use in insects, encompassing 30 different genera. We know that capuchin monkeys have been using stone tools to process food for at least 3000 years ( Falótico et al., 2019) but presumably the use of tools appeared even earlier in invertebrates. Most of the reports concern vertebrates, particularly primates and birds, which can manufacture tools to solve specific tasks ( Hunt and Gray, 2002 Sanz et al., 2013 Auersperg et al., 2014), use multiple tools sequentially ( Martin-Ordas et al., 2012), and choose effective tools based on their functional properties ( Visalberghi et al., 2009).
Tool use is defined as "the external employment of an unattached or manipulable attached environmental object to alter more efficiently the form, position, or condition of another object, another organism, or the user itself, when the user holds and directly manipulates the tool during or prior to use and is responsible for the proper and effective orientation of the tool" ( Shumaker et al., 2011, p. Tool use is a widespread phenomenon within the animal kingdom ( Shumaker et al., 2011 Sanz et al., 2013) and new examples of animal tool use are regularly discovered, such as recently in pigs ( Root-Bernstein et al., 2019) and seabirds ( Fayet et al., 2020).
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