Antipredator benefits of group living in colonial web-building spiders: the .early warning' effect

The Mexican colonial web-building spider Metepeira incrassata is frequently attacked by predatory wasps from a number of families. Previous studies have shown that wasps often attack more than one spider during a ‘run’ on a colony, but capture success declines as colony size increases, and that spiders in the central core of the colony have lower risk (Rayor & Uetz 1990, Behavioral Ecology and Sociobiology, 27, 77–85; Uetz & Hieber 1994, Behavioral Ecology, 5, 326–333). We used data from direct observation of attacks and field experiments to test the hypothesis that colonial web-building spiders benefit from ‘early warning’ of predator approach through vibrations in the colony web. Analysis of 135 naturally occurring wasp attack ‘runs’ (attacks on 454 spiders) showed that the per-attack run and per-spider capture success of wasps decreased significantly with increased spider colony size. Spider defensive and evasive behaviours observed in a subset of these attacks varied with the attack sequence. Evasive responses by spiders were more frequent later in the attack, suggesting advance warning of predator approach. Experiments using a predator-simulating vibration source demonstrated that mean reaction distance of spiders increased with increasing colony size. Adult female spiders in core positions reacted at greater distances than those on the periphery, but immature spiders, whose capture risk is lower, showed no difference. Behaviour of spiders during simulated attacks was similar to observed encounters with wasps: evasive responses were more frequent and response latency was shorter in spiders attacked later in the sequence, and in many cases, spiders took evasive action prior to any contact with the stimulus. Additional experiments testing isolated cues (web contact, airborne vibration, web-borne vibration) suggest spiders respond to web-borne vibrations generated by predators and evasive behaviours of other spiders. Together, these results support the ‘early warning’ hypothesis of antipredator benefits for colonial web-building spiders.