Spider orientation and hub position in orb webs

Orb-web building spiders (Araneae: Araneoidea, Uloboridae) can be considered as territorial central place foragers. In territorial central place foragers, the optimal foraging arena is circular, with the forager sitting in its centre. In orb webs, the spider's orientation (head up or head down) whilst waiting for prey on the hub of its web and the downwards-upwards asymmetry of its running speeds are the probable causes for the observed deviation of the hub from the web's centre. Here, we present an analytical model and a more refined simulation model to analyse the relationships amongst the spider's running speeds, its orientation whilst waiting for prey and the vertical asymmetry of orb webs. The results of our models suggest that (a) waiting for prey head down is generally favourable because it allows the spider to reach the prey in its web on average quicker than spiders waiting head up, (b) the downwards-upwards running speed asymmetry, together with the head-down orientation of most spiders, are likely causes for the observed vertical asymmetry of orb webs, (c) waiting head up can be advantageous for spiders whose downwards-upwards running speed asymmetry is small and who experience high prey tumbling rates and (d) spiders waiting head up should place their hub lower than similar spiders waiting head dow