Artificial light and sun compass orientation in the sandhopper Talitrus saltator (Crustacea, Amphipoda)

Experiments on compass orientation under artificial light were conducted with adult individuals of Talitrus saltator. The aim was to reproduce in the laboratory an orientation based on the sun compass corresponding to that recorded in conditions of the true sun and sky. This was obtained by the creation within an opaque Plexiglas dome of a scenario that permitted variation of the brightness of the artificial sky and sun. The results show that it is possible to obtain sun compass orientation corresponding to the natural situation even in an artificial environment. It can be concluded that sandhoppers identify an artificial light source as the sun if the artificial sky is also illuminated and if the intensities of the artificial sun and sky exceed certain threshold values (1.13 and 10 μW cm⁻², respectively). The results of other experiments under the natural blue sky with an artificial sun and with the real sun attenuated are discussed. Accepted: 23 May 1997     

Moon orientation in adult and young sandhoppers under artificial light

Our experiments, carried out at night and during the day on adults and laboratory-born young of the sandhopper Talitrus saltator, deal with the identification and use of the moon as an orientating factor. Sandhoppers were released in an apparatus (a Plexiglas dome) that produced a scenario similar to the natural one (with artificial sky, moon or sun illuminated at different intensities).When tested at night, the adult and young sandhoppers used the artificial moon like the natural one, independently of the intensity of illumination of the artificial sky and moon. In other words, sandhoppers tested at night always identified the artificial moon as the moon and never as the sun. In daytime releases, the seaward orientation failed at low intensities of artificial sky and sun illumination (3.07 and 1.55 microW cm2, respectively), whereas the sun compass was used effectively at higher levels of artificial sun and sky illumination. The innate ability of moon compass orientation in inexpert young sandhoppers was demonstrated even under artificial light.     

The role of skylight polarization in the orientation of a day-migrating bird species

To assess the role of skylight polarization in the orientation system of a day-migrating bird, Yellow-faced Honeyeaters (Lichenostomus chrysops, Meliphagidae) were tested in funnel cages for their directional preferences. In control tests in the natural local geomagnetic field under the clear natural sky, they preferred their normal migratory course. Manipulations of the e-vector by depolarizing the skylight or rotating the axis of polarization failed to affect the orientation as long as the natural geomagnetic field was present. When deprived of magnetic information, the birds continued in their normal migratory direction as long as they had access to information from the natural sky, or when either the sun or polarized light was available. However, when sun was hidden by clouds, depolarizers caused disorientation. — These findings indicate that polarized skylight can be used for orientation when no other known cues are available. However in the hierarchy of cues of this species, the polarization pattern clearly ranks lower than information from the geomagnetic field.     

Orientation to shorelines by the supratidal amphipod Talorchestia longicornis: Wavelength specific behavior during sun compass orientation

If released in water or on sand the supratidal amphipod Talorchestia longicornis Say amphipods moves in the onshore direction. The present study was designed to determine whether this species uses the sun as a cue for orientation and if so, which visual pigment in the compound eyes is involved. When tested in an apparatus with a view of only the sun and sky amphipods were disoriented when the sun was obscured by clouds. However, when the sun was visible, they oriented in the onshore direction of their home beach in both water and air during both the morning and afternoon. Resetting the time of their circadian rhythm in activity with either an altered light:dark or diel temperature cycle also reset the chronometric mechanism associated with sun compass. orientation. T. longicornis has two visual pigments with absorption maxima near 420 nm and 520 nm. Only the 420 nm pigment is used for sun compass orientation, which may be an adaptation for increasing the contrast between the sun and background scattered skylight or for detecting the radiance distribution of skylight. Irradiating the 520 nm absorbing pigment alone induced positive phototaxis to the sun but not onshore orientation. Thus, T. longicornis shows wavelength specific behavior by using only one of its visual pigments for sun compass orientation.     

Orientation in a desert lizard (Uma notata): time-compensated compass movement and polarotaxis

Summary The diurnal escape response of fringetoed lizards (Uma notata) startled by predators demonstrates clear directional orientation not likely to depend on local landmarks in the shifting sands of their desert environment. Evidence that celestial orientation is involved in this behavior has been sought in the present experiments by testing the effects of (1) phase shifting the animal's internal clock by 6 h and (2) by training the lizards to seek shelter while exposed to natural polarization patterns. In the first case, 90° shifts in escape direction were demonstrated in outdoor tests, as expected if a time-compensated sun or sky polarized light compass is involved. In the second instance, significant bimodale-vector dependent orientation was found under an overhead polarizing light filter but this was only evident when the response data were transposed to match the zenithe-vector rotation dependent on the sun's apparent movement through the sky. This extends to reptiles the capacity to utilize overheade-vector directions as a time-compensated sky compass. The sensory site of this discrimination and the relative roles of sun and sky polarization in nature remain to be discovered.     

Moon and sun compasses in sandhoppers rely on two separate chronometric mechanisms

The relationship between the chronometric system of compensation for the apparent movement of the sun and that for the moon has been the subject of several, never proven, hypotheses. Our studies on sandhoppers have demonstrated that the chronometric mechanism of the moon compass is separate from that of the sun compass. They show (i) that a period of seven days in constant darkness has no influence on the capacity for orientation, either solar or lunar, and indicates the presence of one or more continuously operating timing mechanisms; (ii) that two different shifts in the light–dark phase have no effect on the chronometric mechanism of lunar orientation, but they do affect that of solar orientation; and (iii) that exposure to an artificial moon delayed by seven days with respect to the natural cycle causes the expected change in the mean direction of individuals tested under the natural moon, but not of those tested under the sun.     

Talorchestia tricornuta Shoemaker (Amphipoda,Talitridae) from Sandy Shores of Gabon: Compass Mechanisms of Orientation

We studied the capacity for zonal recovery in an equatorial sandhopper, Talorchestia tricornuta, inhabiting the sandy beaches of equatorial West Africa. These beaches are often narrow and backed by an evident landscape. The sky usually has heavy cloud cover. The experiments were performed both in a confined environment and in the field, in full sun or an overcast sky, with the magnetic field natural, deviated or absent T. tricornuta was able to assume the correct direction using the sun and magnetic field as orienting factors. Moreover, the releases in the field with the natural landscape visible indicate that the landscape may constitute a further important orienting factor in zonal recovery.     

The Local Optical Factor as an Orientation Cue for the Sandhopper Talitrus Saltator Montagu (Amphipoda,Talitridae)

The aim of the present study was to assess the influence of the local optical factor (i.e., inhomogeneity in spectral composition between the hemidomes of sky over the sea and over land at the beach) on the ability of zonal orientation of the amphipod Talitrus saltator. The experiments were conducted in the laboratory using coloured filters and an apparatus that provides for orientation in artifcial conditions comparable to that observed under natural sun and sky. The results show that T. saltator can effectively use the inhomogeneity between the two hemidomes, but only if the colour distance between them is much greater than that found in the wild. This makes the possibility of perception - and thus use - of the local optical factor rather unlikely in natural conditions.     

The Magnetic Field as a Reference System for Genetically Encoded Migratory Direction in Pied Flycatchers (Ficedula hypoleuca Pallas)

To see whether the migratory orientation of pied flycatchers (Ficedula hypoleuca Pallas) is genetically encoded with respect to the earth magnetic field a group of young birds was hand-raised. They were thus prevented from ever experiencing the sky. The birds were tested in autumn 1980 and 1981 in the local geomagnetic field (Fig. 1) and in three artificial fields (Fig. 2a-c). The results show that their magnetic compass matures independent of any experience with the sky and contains sufficient information for the birds to orient toward their migratory direction.     

Orientation by magnetic field in leaf-cutter ants, Atta colombica (Hymenoptera: Formicidae)

Leaf‐cutter ants (Atta colombica) use trail following to travel between foraging sites and the home nest. However, this combination of pheromone and visual cues is likely to be complemented by a directional reference system such as a compass, used not only when foraging but also during colony formation, where foraging trails degrade or where ants become displaced. One candidate system is the magnetic polarity compass. We tested the orientation of leaf‐cutter ants under a magnetic field of reversed‐polarity, with the prediction that the ants would show 180° deflection compared with control ants in an unchanged geomagnetic field. When the sun's disc was unobstructed by clouds, orientation was the same as that of control ants, implying that magnetic cues were not used to orient. However, when the sky was overcast, ants in the experimental treatment significantly shifted their mean orientation both in comparison with controls and reversed‐polarity ants under the sun. Although a total reversal in orientation was not induced, the results demonstrate that Atta respond to magnetic reversal in the absence of sunlight cues, and suggest a role for magnetic cues in determining direction during orientation.     

Hierarchy of sun,beach slope,and landmarks as cues for Y-axis orientation of the supratidal amphipod Talorchestia longicornis (Say)

The semi-terrestrial amphipod Talorchestia longicornis (Say) undergoes Y-axis orientation and has a hierarchy among orientation cues. A previous study found that they used sun compass orientation and moved in the onshore direction of the home beach in both air and water. The present study determined whether this species could also use local landmarks and beach slope as orientation cues. They oriented upslope in simulated darkness in the laboratory on both dry and wet sand with threshold slopes of 2° and 4°, respectively. When tested outside in an arena in air on wet sand, they were disoriented when sun, slope, and landmarks were absent as cues. If presented with single cues, they moved upslope, toward landmarks and in the up-beach direction of the home beach during sun compass orientation. Using paired cues, sun was dominant over slope and landmarks, while slope was dominant over landmarks. In the presence of all three cues, amphipods displayed sun compass orientation in all test combinations except when slope and landmarks were paired together against the sun, which evoked a bimodal response. Thus, the hierarchy of cues for up-beach movement of T. longicornis during Y-axis orientation is the sun, then the slope, and finally the landmarks.     

Mass cultivation of Nannochloropsis sp. in annular reactors

A study was made on the mass cultivation of Nannochloropsis sp. in newly designed annular reactors operated under natural, artificial or combined illumination. The annular reactor consists of two 2-m-high Plexiglas cylinders of different diameter placed vertically one inside the other so as to form an annular culture chamber. Artificial illumination is supplied by lamps placed inside the inner cylinder. Two annular reactors of different diameter (50 and 91 cm), light path (4.5 and 3.0 cm) and illuminated surface area (5.3 and 9.3 m²) were experimented with. The effect of two different artificial light sources (fluorescent tubes and metal halide lamps) on culture productivity was investigated in both systems. The highest productivity on a per reactor basis (about 34 g (d. wt) reactor^–1 24 h^–1) was achieved with the larger reactor illuminated by a 400-W metal halide lamp. From February to May a 91-cm reactor illuminated only with natural light was operated in parallel with a 91-cm reactor subjected to combined illumination. Under natural illumination productivity increased from 16.6 g (d. wt) reactor^–1 d^–1 in February to 34.1 g (d. wt) reactor^–1 d^–1 in May. Under combined illumination productivity was 41.3 g (d. wt) reactor^–1 d^–1 in February and increased up to 48.3 g (d. wt) reactor^–1 d^–1 in May. Although the culture exposed to combined illumination always attained higher yields, the productivity gap between the two cultures decreased gradually along the season as solar radiation and minimum night temperatures increased. A 1200-L plant made of ten 50-cm annular reactors was set up and operated for two years with combined illumination yielding an average of 270 g of dry Nannochloropsis sp. biomass per day. More than 2000 L of concentrate suspension (50 g (d. wt) L^–1) of Nannochloropsis sp. were produced and successfully used by fish hatcheries as live feed for rotifers and for rearing seabream larvae with the green-water technique. This study indicates that the annular reactor can be profitably used for long-term cultivation of Nannochloropsis in temperate climates. Besides reliability and ease of operation, the main advantage of the system is that it can be used under natural illumination, yet artificial light can be also supplied to maintain high productivity levels in winter or on cloudy days.     

Microclimatic differences between and within canopy gaps in a temperate rainforest

The microclimate in the understorey and in two canopy gaps of different size (249 m² and 12.6 m²) within a temperate rainforest in northern New Zealand were characterised using net all-wave radiation (Q ^*), vapour pressure deficit (VPD), vertical air temperature gradient (VTG), and maximum and minimum air temperatures. Hemispherical fisheye photographs were used to define the exposure to the sky at each microclimate recording site both within and between gaps, and in the understorey. Under condiions of clear skies during the summer, with the sun close to its maximum altitude, the large gap centre had a mean daytime (0700–1700 hours)Q ^* of 452 W/m². This value was similar to that estimated for an open field nearby, but three times greater thanQ ^* at the centre of the small gap, and six times greater than in the understorey. The east and west sides of both gaps had higher values ofQ ^* than the north and south sides. BothQ ^* and mean daily maximum temperature were strongly correlated with the amount of sky exposure. VPD and VTG were higher in gaps than in the understorey, with maximum values for both variables occurring at the centre of the large gap.     

Islands within islands: two montane palaeo‐endemic birds impacted by recent anthropogenic fragmentation

Anthropogenic habitat fragmentation of species that live in naturally patchy metapopulations such as mountaintops or sky islands experiences two levels of patchiness. Effects of such multilevel patchiness on species have rarely been examined. Metapopulation theory suggests that patchy habitats could have varied impacts on persistence, dependent on differential migration. It is not known whether montane endemic species, evolutionarily adapted to natural patchiness, are able to disperse between anthropogenic fragments at similar spatial scales as natural patches. We investigated historic and contemporary gene flow between natural and anthropogenic patches across the distribution range of a Western Ghats sky‐island‐endemic bird species complex. Data from 14 microsatellites for 218 individuals detected major genetic structuring by deep valleys, including one hitherto undescribed barrier. As expected, we found strong effects of historic genetic differentiation across natural patches, but not across anthropogenic fragments. Contrastingly, contemporary differentiation (D_PS) was higher relative to historic differentiation (F_ST) in anthropogenic fragments, despite the species’ ability to historically traverse shallow valleys. Simulations of recent isolation resulted in high D_PS/F_ST values, confirming recent isolation in Western Ghats anthropogenic fragments and also suggesting that this ratio can be used to identifying recent fragmentation in the context of historic connectedness. We suggest that in this landscape, in addition to natural patchiness affecting population connectivity, anthropogenic fragmentation additionally impacts connectivity, making anthropogenic fragments akin to islands within natural islands of montane habitat, a pattern that may be recovered in other sky‐island systems.     

Orientation of the sandhopperTalitrus saltator (Amphipoda,Talitridae) living on dynamic sandy shores

Summary The littoral sandhopperTalitrus saltator Montagu maintains its position along the beach mainly by means of an innate solar orientation. Different populations show different directional tendencies depending on the direction of the sealand axis. Experiments were designed which aimed to disentangle the innate and acquired components of the escape orientation. Therefore, the solar orientation of experienced adults living on shores subject to fluctuations of the shoreline and of their laboratory-born (inexperienced) offspring were compared. The innate sun compass orientation shown by the young is modified by learning during life under natural conditions, depending on the variability of the shore. Results showed: (1) an improvement in the solar orientation in individuals living on relatively constant shores; (2) a loss of precision in individuals living on highly dynamic shores; (3) a change to a new ecologically efficient escape direction in individuals living on a relatively stable lagoon shore.Abbreviation TED theoretical escape direction     

Locomotor activity rhythm and sun compass orientation in the sandhopper <Emphasis Type="Italic">Talitrus </Emphasis><Emphasis Type="Italic">saltator</Emphasis> are related

The sandhopper Talitrus saltator has an endogenous activity rhythm with a circadian periodicity. It is well known for its ability to compensate for the apparent movement of the sun during its migrations along the sea–land axis of the beach. Both chronometric mechanisms are entrained by the natural LD photoperiod. Using actographic recordings and tests of solar orientation of individuals kept under an LD 12:12 clock-shifted cycle, after 1–13 days of treatment, we demonstrate that the timing mechanism of activity rhythm and the chronometric mechanism underlying the sun compass are the same.     

Polarized skylight orientation in the desert antCataglyphis

Summary The desert antCataglyphis bicolor is able to use the pattern of polarized light in the sky as compass. By confronting the ant to single spots of artificially and naturally polarized light it is shown howCataglyphis uses the polarization pattern.When exposed to a horizontal e-vector,Cataglyphis was always oriented correctly. Orientation errors occurred, however, when other e-vector directions were presented. This indicates that the e-vector positions assumed by the ant do not coincide with the e-vector positions actually realized in the sky. From this it is concluded thatCataglyphis has no detailed knowledge of the actual azimuthal positions of the e-vectors. Instead, it is relying on a simplified celestial map of the polarization patterns in the sky (Fig. 7).Usually, the ant did not confuse celestial spots with identical e-vector directions. Even at sunset when the polarization pattern is completely ambiguous, correct orientation occurred. This suggests that the ant uses additional celestial cues such as the degree of polarization, the color or the intensity to find its way home when the sun is obscured.     

Light energy partitioning in photosystems I and II during development of Nothofagus nitida growing under different light environments in the Chilean evergreen temperate rain forest

Nothofagus nitida (Phil.) Krasser (Nothofagaceae) regenerates under the shade. Nonetheless, older seedlings are commonly found at full sun. We tested the hypothesis that light capture and photochemical and non-photochemical energy dissipation of both photosystems PSI and PSII adjust with ontogeny and brighter environment. Light energy partitioning in both photosystems was studied in seedlings of different developmental stages (small 9.7 cm, tall 36 cm) under contrasting light environments (8–200 and 1,800–2,043 μmol photons m⁻² s⁻¹) in the Chilean evergreen temperate forest. Higher A _max, dark respiration, and light compensation and saturation points in sun seedlings of both developmental stages were accompanied by higher rates of electron transport. These seedlings also showed a high fraction of open PSII reaction centres and similar non-photochemical quenching at high-light in both photosystems, showing no effect of developmental stage in these parameters. Conversely, light capture, total thermal dissipation after photoinhibition, active down-regulation of antenna efficiency, and state transitions were higher in smaller seedlings than in taller ones. These changes maintain photostasis, preventing photodamage, while favouring a more oxidized quinone pool. There is an independent effect of seedling development and light acclimation on this transition from shade to sun during early ontogeny. This transition reflects short-term responses of the photosynthetic apparatus to light and longer term responses that depend on seedling developmental stage.     

Orientation at night: an innate moon compass in sandhoppers (Amphipoda: Talitridae)

The supralittoral amphipod Talitrus saltator is well known for its capacity for astronomical orientation using the sun and moon as compasses. It has also been demonstrated that the sun compass is innate in this species. In our experiments, we released inexpert (naive) young born in the laboratory into a confined environment under the full moon and in the absence of the horizontal component of the magnetic field. They were allowed to see the natural sky and the moon only at the moment of release. The young individuals were obtained in the laboratory by crossing adult individuals from the same and different populations of sandhoppers. The young from intrapopulation crosses were well oriented towards the directions corresponding to those of their parents, whereas the young from interpopulation crosses were oriented in an intermediate direction. Therefore, our experiments demonstrate in the sandhopper T. saltator that sea-land moon orientation relies on an innate chronometrically compensated mechanism.     

Oriented swimming at an angle to light in a schooling characid, Moenkhausia dichrowa Kner

Two groups each of approximately 100 Moenkhausia dichroua , a schooling characid, showed a long–lasting, constant–oriented swimming when placed in a light–centred circular channel. This apparatus consists of a 1–m diameter circular channel illuminated by either a central or a peripheral light system, so that the light angle is constant all around the channel. With the central light at a fixed angle, fish swam for several months in one direction and reversed direction at a certain date. When the light angle was increased by 10° every other day between 0° (horizontal) and 90° (vertical), swimming direction was reversed at a particular angle in each experiment. This response to artificial light suggests that this small schooling fish uses the sun as an orientation clue in its seasonal migrations.