Animals use many different mechanisms to navigate in space. The characteristics of the mechanism employed are usually well-suited to the demands of each particular navigational problem. For example, desert ants navigating in a relatively featureless environment use path integration, birds homing or migrating over long distances use compasses of various sorts, salmon returning to their natal stream home on olfactory cues. The study of navigation requires the study of many different taxa confronting different problems. One interesting case involves scatter-hoarding species that use memory to relocate their hidden food. Such animals face the problem of remembering many locations simultaneously. Clark's nutcrackers (Nucifraga columbiana) are an excellent example, and this paper considers their possible use of multiple bearings from landmarks.

Predicted flight trajectories differ depending on which orientation cues are used by migrating birds. Results from radar and satellite tracking of migrating birds can be used to test which of the predicted trajectories shows the best fit with observed flight routes, supporting the use of the associated orientation mechanism. Radar studies of bird migration at the Northeast Passage and the Northwest Passage support the occurrence of migration along sun compass routes in these polar regions. In contrast, satellite tracking of Brent geese (Branta bernicla) migrating from Iceland across Greenland and from Northwest Europe to Siberia show routes that conform most closely with geographic loxodromes, but which are also profoundly influenced by large-scale topography. These evaluations are discussed in relation to the adaptive values of different routes in different parts of the world. Sun compass routes are favourable mainly for east-west migration at high latitudes. For east-west migration at mid and high latitudes magnetic loxodromes are more favourable than geographic loxodromes in certain regions while the reverse holds in other regions. The geometry of migration routes, as recorded by radar and satellite tracking, may be important for understanding the evolution of the complexity of birds' orientation systems, and for providing clues about the orientation mechanisms guiding the birds on their global journeys.

Birds can accurately navigate over hundreds to thousands of kilometres, and use celestial and magnetic compass senses to orient their flight. How birds determine their location in order to select the correct homeward bearing (map sense) remains controversial, and has been attributed to their olfactory or magnetic senses. Pigeons can hear infrasound down to 0·05 Hz, and an acoustic avian map is proposed consisting of infrasonic cues radiated from steep-sided topographic features. The source of these infrasonic signals is microseisms continuously generated by interfering oceanic waves. Atmospheric processes affecting the infrasonic map cues can explain perplexing experimental results from pigeon releases. Moreover, four recent disrupted pigeon races in Europe and the north-eastern USA intersected infrasonic shock waves from the Concorde supersonic transport. Having an acoustic map might also allow clock-shifted birds to test their homeward progress and select between their magnetic and solar compasses.

Migrating pierid butterflies use a sun compass in order to head in a preferred flight direction. When blown off-course by the wind, butterflies adjust their heading to compensate for cross-track wind drift. They are capable of drift compensation over water when and where landmarks are visible on the shore. They are also capable of a more inaccurate form of drift compensation over the sea when and where landmarks are not visible.

Albatrosses and sea turtles are known to perform extremely long-distance journeys between disparate feeding areas and breeding sites located on small, isolated, oceanic islands or at specific coastal sites. These oceanic journeys, performed mainly over or through apparently featureless mediums, indicate impressive navigational abilities, and the sensory mechanisms used are still largely unknown. This research used three different approaches to investigate whether bi-coordinate navigation based on magnetic field gradients is likely to explain the navigational performance of wandering albatrosses in the South Atlantic and Indian Oceans and of green turtles breeding on Ascension Island in the South Atlantic Ocean. The possibility that magnetic field parameters can potentially be used in a bi-coordinate magnetic map by wandering albatrosses in their foraging area was investigated by analysing satellite telemetry data published in the literature. The possibilities for using bi-coordinate magnetic navigation varied widely between different areas of the Southern Oceans, indicating that a common mechanism, based on a bi-coordinate geomagnetic map alone, was unlikely for navigation in these areas. In the second approach, satellite telemetry was used to investigate whether Ascension Island green turtles use magnetic information for navigation during migration from their breeding island to foraging areas in Brazilian coastal waters. Disturbing magnets were applied to the heads and carapaces of the turtles, but these appeared to have little effect on their ability to navigate. The only possible effect observed was that some of the turtles with magnets attached were heading for foraging areas slightly south of the control turtles along the Brazilian coast. In the third approach, breeding female green turtles were deliberately displaced in the waters around Ascension Island to investigate which cues these turtles might use to locate and return to the island; the results suggested that cues transported by wind might be involved in the final stages of navigation.

This and the following paper were first presented at the RIN01 Conference held in Oxford under the auspices of the Animal Navigation Special Interest Group, April 2001.

Flight paths of homing pigeons were measured with a newly developed recorder based on GPS. The device consists of a GPS receiver board, a logging facility, an antenna, a power supply, a DC-DC converter and a casing. It has a weight of 33 grams and works reliably with a sampling rate of 1 Hz for an operating time of about three hours, providing time-indexed data on geographic positions, ground speed and altitude. The devices are fixed to the birds with a harness, and the data are downloaded when the bird is re-captured. The measured flight paths show many details : for example, initial loops flown immediately after release and large detours flown by some pigeons. Three examples of flight paths are presented from a release site 17·3 km northeast of the home loft in Frankfurt. Mean speed in flight, duration of breaks and total length of the flight path were calculated. The pigeons chose different routes and have different individual tendencies to fly loops over the village close to the release site.

This paper was first presented at the RIN97 Conference held in Oxford under the auspices of the Animal Navigation Special Interest Group, April 1997.

Migrating species may utilise hydrostatic pressure. In the aquatic environment, hydrostatic pressure changes much more rapidly than in air. In shallow water, tidal changes will impose larger percentage changes on organisms than those experienced in deep water. Small changes in pressure often cause locomotion (barokinesis) accompanied by orientation to light or gravity, often partially compensating for the equivalent depth change. Until recently, identification of hydrostatic pressure receptors without a gas phase has proved elusive, but it is now known that thread hair receptors in the statocyst of the shore crab Carcinus maenas respond to small changes in hydrostatic pressure. Using a tide machine, the responses of thread hairs to sinusoidally changing pressure cycles have been examined, and this paper reports progress monitoring this receptor and making long-term recordings from hydrostatic pressure sensitive pathways in the crab's nervous system.

This paper, and the following 5 papers, were presented at the RIN93 Conference on “Orientation and Navigation - Birds, Humans and other Animals.” This took place in Oxford in April, and included some 40 papers. Copies of the complete Proceedings, priced £25 for RIN members and £35 for non-members, are available from the Director.

According to comprehensive theories of navigation, animals navigate by using two complementary strategies: (1) dead reckoning informs the subject in a continuous manner on its actual location with respect to an Earthbound or absolute coordinate system; while (2) long-term associations between particular landmarks and specific locations allow the animal to find its way within a familiar environment. If the subject structures familiar space as a system of interconnected places – the so-called ‘cognitive map’ – it may know through dead reckoning where it is located on its map and relate its route-based expectations to the actually perceived scenario of local cues.

A model avian compass is formed if magnetite crystals are included among the statoconia loading some of the hair cells in the vestibular system. The magnetite is used as a null detector to detect the direction in which the magnetic couple acting on the magnetic crystal is zero. The location of the north/south plane and the angle between the directions of the gravitational and the magnetic fields are detected at a single site.

Migratory directions of birds have long been assumed to be at least partly ‘innate’ that is, genetically determined. In recent years much progress has been made toward understanding the genetic basis of several components of the orientation system. An important distinction has to be made between a bird's ‘knowledge’ of which direction to migrate, and its ability to use certain compass systems to find this intended direction. Both processes, establishing an intended direction and the subsequent compass orientation, have a partly genetic basis. The present review will be concerned only with the first of these processes, and the way it allows evolutionary changes of migratory routes to occur.

It is now well established that homing pigeons (Columba livia) use a variety of mechanisms and cues to help them find their way home. It is far more difficult to determine the relative importance attached to such mechanisms and cues by pigeons during homing. As a step towards this quantitative stage in our understanding of homing behaviour we consider if cues contribute significantly to homing speed. In particular we attempt to assess the role of visual landmarks. Using pigeons released from familiar release sites, we demonstrate that birds home faster if they can view their surroundings from inside a Perspex box for 5 minutes prior to being released, compared to birds denied this view. Pigeons released from unfamiliar sites, however, do not show a similar improvement. We suggest that seeing familiar visual landmarks before release allows pigeons to become more certain of their current location with respect to home. Simply viewing the release site is not sufficient to produce a faster homing speed; the pigeons must be able to recognize that the release site is familiar.

Many animals have some kind of ‘home’ –a nest, a burrow, etc. This applies to insects that build nests and provide their larvae with food just as much as to numerous species of vertebrates. Rodents and moles are known to dig burrows; most other mammals, reptiles, amphibians and even many fresh-water fish do not roam around randomly either, but have well-defined home ranges with favourite resting places in particular cavities, under rocks or in dense thickets. Birds build nests for their young and fly out searching for food. All these animals permanently face orientation problems: they have to move swiftly and efficiently within their home range, and they must be capable of returning home when they have left their familiar area while exploring, fleeing, etc.