Numerical discrimination of sets of elements by Archerfish

Debates have arisen as to whether non-human animals actually can learn abstract non-symbolic numerousness or whether they always rely on some continuous physical aspect of the stimuli, covarying with number. Here we investigated archerfish (Toxotes jaculatrix) non-symbolic numerical discrimination with accurate control for co-varying continuous physical stimulus attributes. Archerfish were trained to select one of two groups of black dots (Exp. 1: 3 vs. 6 elements; Exp. 2: 2 vs. 3 elements); these were controlled for several combinations of physical variables (elements’ size, overall area, overall perimeter, density and sparsity), ensuring that only numerical information was available. Generalization tests with novel numerical comparisons (2 vs. 3, 5 vs. 8 and 6 vs. 9 in Exp. 1; 3 vs. 4, 3 vs. 6 in Exp. 2) revealed choice for the largest or smallest numerical group according to the relative number that was rewarded at training. None of the continuous physical variables, including spatia...