Nephila clavipes spiders (Araneae: Nephilidae) keep track of captured prey counts: testing for a sense of numerosity in an orb-weaver

Nephila clavipes golden orb-web spiders accumulate prey larders on their webs and search for them if they are removed from their web. Spiders that lose larger larders (i.e., spiders that lose larders consisting of more prey items) search for longer intervals, indicating that the spiders form memories of the size of the prey larders they have accumulated, and use those memories to regulate recovery efforts when the larders are pilfered. Here, we ask whether the spiders represent prey counts (i.e., numerosity) or a continuous integration of prey quantity (mass) in their memories. We manipulated larder sizes in treatments that varied in either prey size or prey numbers but were equivalent in total prey quantity (mass). We then removed the larders to elicit searching and used the spiders' searching behavior as an assay of their representations in memory. Searching increased with prey quantity (larder size) and did so more steeply with higher prey counts than with single prey of larger sizes. Thus, Nephila spiders seem to track prey quantity in two ways, but to attend more to prey numerosity. We discuss alternatives for continuous accumulator mechanisms that remain to be tested against the numerosity hypothesis, and the evolutionary and adaptive significance of evidence suggestive of numerosity in a sit-and-wait invertebrate predator.