Tactile senses and foraging in birds, with emphasis on kiwi : a thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Ecology at Massey University, Manawatu, New Zealand

Abstract

Probe-foraging birds must often rely on senses other than vision for prey-detection. One such sense is ‘remote touch’: the detection of vibration and pressure cues from prey within the substrate. Remote touch is mediated by the ‘scolopacid-type bill-tip organ’, which consists of a honeycomb of sensory pits in the bill-tip, containing clusters of mechanoreceptors. This organ was originally described in the neognathous shorebird family Scolopacidae, but was recently also discovered in paleognathous kiwi (Apterygidae): an example of convergent/parallel evolution. My aim was to discover how widespread this organ is among birds, compare its anatomy and function in foraging between kiwi and other probe-foraging birds and elucidate in detail the foraging behaviours and senses used by free-living kiwi. Within the thesis I compare the bill-tip organs of kiwi and shorebirds using material from the brown kiwi (Apterygidae: Apteryx mantelli) and bar-tailed godwit (Scolopacidae: Limosa lapponica). I provide the first description of the organ in a third family of birds, the ibises (Threskiornithidae), and give evidence that it may exist in simplified form in a fourth family, Rallidae. The Scolopacidae, Apterygidae, Threskiornithidae and Rallidae are widely separated on the avian phylogenetic tree. This suggests that the evolution of the scolopacid-type bill-tip organ and associated sense is favoured by a probe-foraging lifestyle. Foraging trials confirm the bill-tip organs of brown kiwi and Madagascar crested ibises (Lophotibis cristata urschi) are functional for remote touch. The ibises rely solely on remote touch to find buried prey, whereas brown kiwi use the sense in conjunction with olfaction. Free-living brown kiwi display no obviously visually-guided behaviours, instead using hearing (head-lifting in response to noises audible to the observer), olfaction (odour sensing behaviour, ‘sniffing,’ in the direction of these sounds) and touch. Kiwi tap ahead with their bill-tip when walking and move their facial bristles forward when foraging, forming a ‘net’ on the ground. The bristle follicles in kiwi (and some other insectivorous bird species) are innervated with Herbst corpuscles, suggesting tactile function. Female kiwi probe on average 30% deeper than males and juveniles, but there are no other differences in foraging behaviour between the sexes.