Short-range Homing in Fiddler Crabs (Ocypodidae, Genus Uca): A Homing Mechanism not Based on Local Visual Landmarks

This field study investigated the spatial strategies and homing ability of two East African fiddler crabs, Uca lactea annulipes and Uca vocans hesperiae, using various experimental procedures. A clear-cut spatial segregation between the two species was observed, with U. l. annulipes occupying areas even a few centimetres above U. v. hesperiae . Females of both species were extremely faithful to their holes while the males exhibited behavioural differences. U. l. annulipes males were quite faithful to their holes; in contrast, U. v. hesperiae males, during a single low tide, visited several holes which they did not defend. Two displacement experiments were conducted in order to identify the stimuli that fiddler crabs use in their homing. In the 'dislocation experiment', the crabs were moved (by hand) 1 or 2 m away from their refuge and released. None of these crabs returned to its burrow and, moreover, the initial directions they followed were not homeward orientated. In the 'translation experiment', the crabs were passively translated while actively feeding. When the crabs attempted to go 'home' after the translation, they darted along a straight path that led them to points that would have corresponded to their homes if the translation had not taken place. After this first response crabs then commenced searching strategies that led half of them home. Short-range homing seems to depend strongly on information actively gathered during the outward journey, other than local visual landmarks.     

Homing in the swimming crab Thalamita crenata: a mechanism based on underwater landmark memory

We investigated whether Thalamita crenata, a swimming crab found on the East African intertidal flats, uses landmarks to locate its refuges. We modified the visual panorama of an intertidal flat, using conspicuous movable objects, and conducted homing trials with the local population of swimming crabs. In the first set of trials, after being moved away from their dens, the crabs were immediately able to find other known dens, using direct shortcut paths. In the second set of trials we moved all the artificial landmarks surrounding a crab's den 5 m away and then displaced the crab itself. The crabs made initial navigational errors in accordance with the new position of the landmarks; this shows that they oriented themselves by remembering the landmarks. We then repositioned the landmarks and released the crabs far from the familiar area, on a similar flat. This time the crabs could rely only on the artificial landmarks; they used this information and reached the point where home should have been according to the landmarks. Thus, T. crenata showed good spatial knowledge, based on the storage of landmark memories. This orienting mechanism is much more flexible and complex than those of other crabs and is comparable to the route-based memory of honeybees, Apis mellifera. Copyright 2000 The Association for the Study of Animal Behaviour.     

The role of geomagnetic cues in green turtle open sea navigation

Background

Laboratory and field experiments have provided evidence that sea turtles use geomagnetic cues to navigate in the open sea. For instance, green turtles (Chelonia mydas) displaced 100 km away from their nesting site were impaired in returning home when carrying a strong magnet glued on the head. However, the actual role of geomagnetic cues remains unclear, since magnetically treated green turtles can perform large scale (>2000 km) post-nesting migrations no differently from controls.

Methodology/Principal Findings

In the present homing experiment, 24 green turtles were displaced 200 km away from their nesting site on an oceanic island, and tracked, for the first time in this type of experiment, with Global Positioning System (GPS), which is able to provide much more frequent and accurate locations than previously used tracking methods. Eight turtles were magnetically treated for 24–48 h on the nesting beach prior to displacement, and another eight turtles had a magnet glued on the head at the release site. The last eight turtles were used as controls. Detailed analyses of water masses-related (i.e., current-corrected) homing paths showed that magnetically treated turtles were able to navigate toward their nesting site as efficiently as controls, but those carrying magnets were significantly impaired once they arrived within 50 km of home.

Conclusions/Significance

While green turtles do not seem to need geomagnetic cues to navigate far from the goal, these cues become necessary when turtles get closer to home. As the very last part of the homing trip (within a few kilometers of home) likely depends on non-magnetic cues, our results suggest that magnetic cues play a key role in sea turtle navigation at an intermediate scale by bridging the gap between large and small scale navigational processes, which both appear to depend on non-magnetic cues.     

Homing in fiddler crabs (Uca lactea annulipes and Uca vomeris : Ocypodidae)

Fiddler crabs emerge from burrows on intertidal sand- and mudflats to feed during low tide. In the species studied here (Uca lactea annulipes, Uca vomeris) a crab normally wanders no more than about 1 m away from its burrow and, when frightened, dashes back along a straight line to take cover. Feeding crabs tend to move sideways, without changing orientation, along paths radiating from the burrow. When they move along circumferential paths they adjust their orientation so that one side continues to point towards the burrow. The crabs do not need to see the burrow in order to stay aligned with the home vector, and they are not misled by a dummy hole close to their own burrow unless they have come to within about 10 cm of it. The home runs of crabs end within a few centimeters of a burrow that is covered with a sheet of sandpaper and then give way to search runs, centred upon a position slightly short of the burrow location. Feeding crabs can be displaced on sandpapers and their subsequent home runs end at a position where the burrow would be, had there been no displacement. Landmarks close to the burrow do not influence the home runs of displaced crabs. Crabs that are rotated on a sheet of sandpaper, counter-turn to keep their original orientation constant. Fiddler crabs thus employ path integration with external compass information and close range visual guidance for homing. Accepted: 11 May 1998     

Mechanisms of visually mediated site recognition by the homing pigeon

The recognition of familiar areas by homing pigeons, Columba livia, is now known to depend at least in part on visual cues. Birds allowed a 5-min preview of the surrounding landscape prior to release home faster than those denied access to such cues, suggesting that recognition is visually mediated. We examined this phenomenon further by asking how memory generated through prior experience with a site is used in recognition. We provided a group of homing pigeons with training experience in which they viewed, through a single transparent vertical face of an otherwise opaque release box, an approximately 140° segment of the landscape for 5 min before release. Training previews were always given from consistent locations and orientations within a release site. Test releases that followed were used to ascertain whether subsequent site recognition necessitated previewing from this already familiar angle and distance, or whether the skill could be extended to novel, nonoverlapping lateral views of the landscape. The results suggested that homing performance was better after presentation of the view recapitulating that seen during training than after showing a novel alternative view. Pigeons may have been using a template of the arrangement of familiar landmarks around the release site as the cue for recognition. The effect, however, disappeared after repeated training, suggesting that repeated release from a site may allow for more extensive visual survey and the memorization of detailed features of the landscape or the extraction of site-specific general features that aid recognition even when a novel view is presented. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Long-distance Homing Ability in Dasypoda altercator (Hymenoptera,Melittidae)

Homing rates and initial orientations after release from different directions were investigated in the solitary bee Dasypoda altercator. Homing rates and the proportion of individuals returning on the day of release declined with distance, implying that homing from greater distances is not based solely on the use of landmarks. A long duration of return (often not on the same day) and high efficiency of homing (four bees out of 10 returned from a 4 km distance over the lake), as well as an increase in average return speed ? 10 m/min) with distance of bees homing on the same day suggest that homing is also not based upon radial scatter as the sole tactic. This is supported by the observation that the initial orientations of the bees were not equally probable. Departures towards the sun greatly exceeded those away from the sun and the bees also tended to depart in cardinal geographical (or geomagnetic) directions, preferring meridional directions (especially southward) to parallel ones (of which east was preferred). Departure directions did not depend on wind direction but did depend, to some extent, on the landscape features of release sites. However, bees neither tended to depart in the direction of the nest, nor did homing success correlate with the direction of departure in relation to that of the nest.     

Trail‐following and trail‐searching behaviour in homing of the intertidal gastropod mollusc,Onchidium verruculatum

When uncovered by the tide, Onchidium verruculatum leaves its place of hiding, grazes on the rock surface, and thèn returns to its own home. Onchidium will follow its own mucus trail, most frequently towards the origin. This behaviour explains homing when the outward path is partly or completely retraced. Sometimes, however, homing occurs without contact with the outward trail. It was observed that trails were followed more closely at low humidities than at high humidities. No evidence was found for any method of orientation other than detection of chemical trials: when displaced onto sand Onchidium tend simply to crawl in a spiral until they contact their own trail. In addition to being a safe retreat, the home provides a fixed starting point to feeding excursions : this may facilitate re‐location of preferred feeding areas.     

The Orientational Consequences of Flocking Behaviour in Homing Pigeons,Columba livia

Two experiments are described which investigate the orientational consequences of flocking in homing pigeons Columba livia. Previous experiments have shown that homing pigeons placed inside a clear-sided release box for 5 min before release from a familiar site have enhanced ground homing speed compared with those placed in an opaque-sided box. It is assumed that previewing the surrounding landscape allows for faster homing since a bird denied this information must accumulate the knowledge on release. In experiment 1, using the same technique developed in these experiments but releasing the birds in pairs we showed that within familiar areas, homing pigeons can exploit a partner that has acquired more information, allowing them to home more quickly. In experiment 2 we attempted to test three potential strategies which may occur during homing flights. The results do not conclusively distinguish between these three mechanisms but suggest that orientation of the pairs of birds is most likely to have resulted from a compromise of individual tendencies, or from following the best homer, but not from following a ‘governing leader’. The consequence of these mechanisms is discussed.     

Homing Experiments in the Tree-climbing Crab Sesarma leptosoma (Decapoda,Grapsidae)

Sesarma leptosoma an East African mangrove-dwelling crab, migrates twice a day from a system of known dens among the roots to well-defined feeding areas in the branches of trees, reaching 15 m high. Field experiments were performed to test whether chemical or visual cues are involved in the orientation and homing of this species to reach their feeding areas. Manipulation of the substratum at branch junctions, in order to alter possible chemical cues, did not affect homing ability in S. leptosoma. Moreover, crabs trained to cross an asymmetrical artificial wooden fork could still follow their preferred directions after (1) the fork branches had been switched, (2) the whole fork had been rotated around the trunk, resulting in a right-left inversion, and (3) the inversion of two wide black and white screens hiding most of the canopy from view of the climbing crabs. These results suggest that S. leptosoma may not rely on reference systems such as chemical trail-following and chemical or visual cues from the substratum, but probably depend on complex visual information from the surroundings trunks and/or from the sun's position integrated with junction sequence memory.     

When Does Previewing the Landscape Affect Pigeon Homing?

The importance of visual landmarks during homing in pigeons (Columba livia) remains a contentious issue. Three experiments which explore the role of visual landmarks at release sites are reported here. The effects of releasing homing pigeons after a 5-minute period in either a clear or an opaque sided release box were investigated. In the clear sided box pigeons were able to observe local surroundings at a release site, but this view was obstructed in the opaque sided box. In experiment 1 pigeons were released from familiar locations close to home (between 2 and 5.6 km): being unable to view landmarks prior to release significantly slowed homing speeds. In experiment 2 pigeons were released at familiar locations further from home (between 8.4 and 10 km): being unable to view landmarks prior to release did not significantly affect homing speeds. In experiment 3 pigeons were trained to home from distant release sites but were tested at closer, unfamiliar sites located on the likely homing routes used by the pigeons in training. No significant difference in homing speeds were observed when pigeons were released from either the clear or opaque sided box. The significance of these results for understanding the role of visual landmarks within a pigeon's familiar area is discussed.     

Dead reckoning in a small mammal: the evaluation of distance

When hoarding food under infra-red light, golden hamsters Mesocricetus auratus W. return fairly directly from a feeding place to their nest site by evaluating and updating internal signals that they have generated during the previous outward journey to the feeding place. To test more specifically the animals' capacity to evaluate the linear components of the outward journey, the subjects were led from their (cone-shaped) nest to a feeding place along a detour which comprised either 2 (experiment 1) or 5 (experiment 2) segments; adjoining segments were at right angles to each other. In these conditions, the subjects remained significantly oriented towards the nest and therefore were capable of assessing translations as well as rotations during the outward journey. In experiment 3, the nest was removed after the hamsters had started the direct outward journey to the feeding place and the hamsters were rotated during the food uptake. The animals were no longer oriented towards the starting point of their journey, but nonetheless covered, along a fairly straight path, the correct homing distance, and then changed over to a circular search path. These results confirm that mammals can derive the linear components of an outward journey from self-generated signals and therefore are able to judge the homing distance without relying on cues from the environment. For a number of detour outward journeys, our data yield an unexpectedly good fit to Müller and Wehner's (1988) model of dead reckoning in ants. However, this is no longer the case when the outward journey contains an initial loop which brings the subject back to the starting point. These findings are discussed in terms of the biological significance and limitations of an approximate form of path integration.     

Use of local cues in the night-time navigation of the wandering desert spider <Emphasis Type="Italic">Leucorchestris arenicola</Emphasis> (Araneae,Sparassidae)

Adult male Leucorchestris arenicola can walk round-trips of several tens of meters in search of females. Most excursions end with the spiders returning to their burrow. For small animals homing over distances of several meters is theoretically impossible without the aid of external cues. It was investigated, whether the spiders use local cues or they rely solely on global cues. Individually marked male spiders were captured during their excursions and displaced several meters inside an opaque box. Ten out of twelve displaced spiders returned to their burrows. This shows that the male L. arenicola are using local cues during their homing, as the comparatively small displacement distances could not be detected by means of global, e.g. celestial cues. In order to test whether the spiders could be using olfactory guidance, the burrows were displaced by 2 m while the spiders were out on their journeys. In 12 out of 15 experiments, the spiders did not find their burrows. These results show that the burrows do not function as olfactory beacons for the homing spiders.     

Rapid learning of shelter position in an intertidal fish, the shanny Lipophrys pholis L.

The homing ability of an intertidal fish, the shanny Lipophrys pholis , was investigated using two experiments that were based on the shanny's natural propensity to home to a refuge. A displacement experiment demonstrated that the fish were able to accurately locate the previous position of a refuge once the shelter itself had been removed so that it could not be used as a cue to directly signal the goal location. This shows that the shanny can encode information about its familiar surroundings into a spatial map and use this information to home. A second experiment in which the cues internal and external to the experimental tank were put in conflict with one another suggested that the shanny can encode cues that are both intra- and external-tank cues in its representation of space, but that there is individual variation in the type of cues that are used, or memorized.     

Homing Behaviour in Polyergus rufescens Latr. (Hymenoptera,Formicidae)

Homing mechanisms of the European slave-making ant Polyergus rufescens Latr. are investigated by field experiments. The analysis of the behaviour and paths of both homing scouts and raiders after passive displacement showed that: i. Scouts probably home by using a path integration system based on celestial cues; and ii. Displaced raiders do not seem to adopt such a vectorial orientation mechanism. Moreover, we found that passively displaced scouts exhibit a systematic search strategy for the nest after a rectilinear path. By contrast, raiders perform a similar search pattern just after release. Similarities between Cataglyphis and Polyergus homing behaviour are discussed.     

Strong homing does not predict high site fidelity in juvenile reef fishes

After being displaced, juvenile reef fishes are able to return home over large distances. This strong homing behaviour is extraordinary and may allow insights into the longer-term spatial ecology of fish communities. For example, it appears intuitive that strong homing behaviour should be indicative of long-term site fidelity. However, this connection has rarely been tested. We quantified the site fidelity of juvenile fishes of four species after returning home following displacement. Two species, parrotfishes and Pomacentrus moluccensis, showed significantly reduced site fidelity after returning home. On average, they disappeared from their home sites almost 3 d earlier than expected. Mortality or competitive exclusion does not seem to be the main reasons for their disappearance. Rather, we suggest an increased propensity to relocate after encountering alternative reef locations while homing. It appears that some juvenile fishes may have a higher innate spatial flexibility than their strict homing drive suggests.

     

Orientation of anosmatic pigeons

Summary Test releases performed at five symmetrically arranged sites around the loft, at a distance of 78–99 km from it, showed that 1) anosmatic birds transported without alteration of the earth's magnetic field were completely random-oriented, 2) anosmatic birds transported in a container inside which the intensity of the magnetic field was strongly reduced were unable to orientate homewards and mostly departed according to a preferred compass direction, 3) control birds, which could smell, and were transported without alteration of the magnetic field, were homeward oriented and performed better in homing than both experimental groups. The conclusion is that anosmatic birds are unable to detect home direction at unfamiliar sites and that magnetic stimuli perceived during the outward journey are unable to substitute olfactory cues.Abbreviation PCD preferred compass direction Supported by a grant from the Consiglio Nazionale delle Ricerche     

The visual perceptual range of a lizard, <Emphasis Type="Italic">Tiliqua rugosa</Emphasis>

The fragmentation of landscapes produces habitat gaps where the distance between visual landmarks may exceed the perceptual range of a species and impose navigational constraints. We estimated the visual perceptual range of the Australian sleepy lizard, Tiliqua rugosa, by releasing individuals in the centre of a cleared arena in high temperature conditions, with a 0.5-m-high bush placed either 10, 20 or 30 m from the release site. Lizards were more likely to locate those bushes and shelter under them when they were closer, and no lizards found a bush at 30 m. In addition, lizards were less likely to move from the release point when bushes were at 30 m than when they were at the two closer distances. These data suggest that for sleepy lizards the perceptual range for a 0.5-m-high bush is about 20 m. In the uncleared chenopod shrub-land where these lizards live, suitable shelter bushes are an average of 10.5 m from any point in their home range, well within their perceptual range.     

Spatial Orientation in Blattella germanica L. Larvae

The aim of these experiments was to investigate the type of cues used in homing processes by young Blattella germanica L. larvae. Several types of stimuli were tested: path integration with kinesthetic cues and visual orientation with landmark cues. Tests measured the escape direction of larvae from the food box after disturbance. Either type of cue alone, path integration or visual landmarks, was sufficient to allow larvae to orient towards their shelters, but they oriented more precisély when both types of cue were used. When several landmark cues (proximal and distal) were present, their relative angular position seemed important in the orientation process. Macroscopic shapes in the environment appeared to be used as a global image, memorized to reach the shelter.     

Adult fiddler crabs Uca pugnax (Smith) enhance sediment-associated cues for molting of conspecific megalopae

Adult-associated chemical cues can stimulate settlement and metamorphosis of invertebrate larvae into habitats with an enhanced likelihood of juvenile and adult survival. For example, sediments from adult fiddler crab habitat stimulate fiddler crab megalopae to metamorphose (molt) sooner than sediments without adult cues. A similar stimulation of molting occurs after exposure to waterborne chemical cues from adult habitats and to exudates and extracts of adult crabs. We tested whether sediments from habitats without adult Uca pugnax (Smith), which do not stimulate molting of their megalopae, could become stimulatory through brief exposure to adult crabs. Sediments were collected from tidal flats at several distances (∼ 1 m, ∼ 50 m, and ∼ 5.4 km) from adult habitats, and incubated for 24 h with or without adult crabs. Molting rates of laboratory-reared megalopae exposed for 48 h to adult-conditioned sediments were compared to those for untreated controls. Sediments collected in or within 1 m of adult habitat elicited the highest molting rates, and natural sediments from 50 m and 5.4 km had little or no effect on molting. However, incubating sediments collected away from adult habitat with adult crabs produced a higher molting response, and the magnitude of the enhancement increased with distance from adult habitat. Results suggest that the chemical cues that adult crabs release are retained by sediments and consequently stimulate molting of megalopae, regardless of the nature of the sediments themselves. Lack of chemical cues may retard colonization of newly created or heavily disturbed habitats that are otherwise suitable settlement and adult habitat.     

Possible use of environmental gradients in orientation by homing wood mice,Apodemus sylvaticus

51 wood mice, Apodemus sylvaticus, tagged with chemoluminescent bulbs, were visually tracked in a series of night homing experiments. The mice did not go straight towards home, but wandered around the release area and covered distances equal to twice the radial lengths recorded. In the course of their walk, they orientated preferentially towards landscapes which resembled their usual habitat, irrespective of where their actual homeward direction lay. In spite of the randomness of their paths, homing success was good, but detailed analysis of homing time showed that the homing performances of the mice depended on the direction in which they vanished in the course of the observation period. The good fit of observed orientations with those obtained by computer simulation of klinokinesis along an axial gradient, and field observation of a vegetational gradient in the study area, are both consistent with the hypothesis that homing may have resulted from wandering, locally biased by a klinokinetic effect