Homing and "ortsteue" (attachment to place) in Patella L. (Gastropoda, Prosobranchia)

Abstract

It is known that numerous gastropods show homing ability. Many limpets return to well defined sites. Patella L. acquires a home spot also on the smooth glass surfaces of aquarium. Before leaving, the limpet scrapes its home, particularly the anterior part of the footprint. After each return it makes characteristic movements with its shell.When homing Patella nearly always retraces its outward track, at least in part; sometimes it reaches its home over the shortest route possible.Site and tracks are marked by chemical substances. The animal orientates itself by contact chemoreception on the track and by distance chemoreception when near its home. Apparently there also exists a kinesthetic mechanism of orientation based on a measurement of distance on the outward journey.At the home spot Patella always manoeuveres exactly into the same position. Often it has to turn or to glide around slightly. It finally settles down with small movements of its shell.The footprint shows chemical differences in a wide marginal zone. Probably, different chemoreceptors on the sole of the foot are associated to the various regions of the footprint.The chemical differentiation of the footprint has the same orientation-value for all individuals of the same size group. In addition there exists an individual marking, shown e.g. by avoidance reactions of alien animals.The individual specificity of the foot's secretum is particularly marked on the tracks, for each Patella follows its own trail only and thus normally reaches its own home. On an uneven substratum this is the only place where the margin of its shell fits the ground exactly.For establishing a home site and keeping to it, several environmental conditions have to be met. Patella settles only on surfaces overgrown with small algae. In an aquarium it favours the sides most exposed to light. On vertical walls the animal is gravity-oriented. Most limpets are attached head-end down. Patella is able to perceive the distance from the water-line, and may orient itself with respect to fixed objects in the aquarium by means of a "Ferntastsinn". When first choosing a home spot on an uneven substratum, the limpet often adjusts its position by turning movements, so that the contacts between shell and ground are as close as possible. Selective growth at the shell's margin then results in a perfect fit to the irregularities of the ground.All prerequisites for the establishment of a home site are also determining the animal's keeping to its home. If changes are introduced with respect to the fitting of the shell to the ground, to the position of neighbouring objects, to water-level, light, or direction of gravity, the animal may leave its home site and reorient itself in an equivalent new one. The attractiveness of the chemical markings of the old site nevertheless compete with all these conditions for some time.Rest and activity are controlled by light-dark and tidal cycles. - Inter-and intraspecific differences were obtained under experimental conditions. Almost all individuals were preferentially active in darkness. P. caerulea moved at high tide; P. vulgata was active either at low tide only, or both at high and low tide, or - rarely-at high tide only. All animals reacting to tidal rhythms were synchronized even by unnaturally frequent changes in the water-level. The tidal dependence of the activity of individual P. vulgata may perhaps be interpreted as the persistence of an adaptation to drought and submergence acquired at the original habitat. The reactive ability towards high and low tide can be modified. The activity-phase may be stopped prematurely by changing light or tidal conditions. Animals keeping constantly to their homes returned to them at once, whereas others - having no defined home -generally prolonged their excursions. When light and water-level are kept constant the locomotory activity of P. vulgata and P. caerulea has the characteristics of an endogenous tidal rhythm. Changes in water-level soon approximately catch the freerunning periodicity, alteration of light and darkness synchronize it less readily but more precisely.