Jan M. Hemmi

Visually mediated species and neighbour recognition in fiddler crabs (Uca mjoebergi and Uca capricornis)

Mating signals are often directed at numerous senses and provide information about species identity, gender, receptiveness, individual identity and mate quality. Given the diversity of colourful body patterns in invertebrates, surprisingly few studies have examined the role of these visual signals in mate recognition. Here, we demonstrate the use of claw coloration as a species recognition signal in a fiddler crab (Uca mjoebergi). Furthermore, we show that distinct carapace colour patterns in Uca capricornis enable males to discriminate between their female neighbours and unfamiliar females. This is the first empirical evidence of the social importance of colour markings in fiddler crabs and the first example of visually mediated species and neighbour recognition in invertebrates other than insects.

Evidence for spatial aliasing effects in the Y-like cells of the magnocellular visual pathway

Several lines of evidence are provided indicating that our visual percept can be dominated by spatial aliasing for viewing conditions near those needed to see the spatial frequency doubled illusion. The apparent aliasing effect indicates that the underlying sampling array has a density 15-30% of that of M-cells, in agreement with the known proportion of Y-like M-cells (M(y)-cells). The presence of aliasing indicates, that there is a separate irregular array of M(y)-cells, and that their role is to rapidly convey information on retinal gain control to the brain rather than to act primarily as inputs to image motion computation.

Signaling against the Wind: Modifying Motion-Signal Structure in Response to Increased Noise

Animal signals are optimized for particular signaling environments [1-3]. While signaling, senders often choose favorable conditions that ensure reliable detection and transmission [4-8], suggesting that they are sensitive to changes in signal efficacy. Recent evidence has also shown that animals will increase the amplitude or intensity of their acoustic signals at times of increased environmental noise [9-11]. The nature of these adjustments provides important insights into sensory processing. However, only a single piece of correlative evidence for signals defined by movement suggests that visual-signal design depends on ambient motion noise [12]. Here we show experimentally for the first time that animals communicating with movement will adjust their displays when environmental motion noise increases. Surprisingly, under sustained wind conditions, the Australian lizard Amphibolurus muricatus changed the structure and increased the duration of its introductory tail flicking, rather than increasing signaling speed. The way these lizards restructure the alerting component of their movement-based aggressive display in the presence of increased motion noise highlights the challenge we face in understanding motion-detection mechanisms under natural operating conditions.

Polarised skylight and the landmark panorama provide night active bull ants with compass information during route following

SUMMARY Navigating animals are known to use a number of celestial and terrestrial compass cues that allow them to determine and control their direction of travel. Which of the cues dominate appears to depend on their salience. Here we show that night-active bull ants attend to both the pattern of polarised skylight and the landmark panorama in their familiar habitat. When the two directional cues are in conflict, ants choose a compromise direction. However, landmark guidance appears to be the primary mechanism of navigation used by forager ants, with those cues in the direction of heading having the greatest influence on navigation. Different colonies respond to the removal of these cues to different degrees, depending on the directional information provided by the local landmark panorama. Interestingly, other parts of the surrounding panorama also influence foraging speed and accuracy, suggesting that they too play a role in navigation.

Predator avoidance in fiddler crabs: 1. Escape decisions in relation to the risk of predation

The risk of predation is a strong force shaping many aspects of animal behaviour. Early detection and efficient avoidance strategies not only help prey to survive, but also limit the negative impact predation has on other aspects of their lives. The type of antipredator strategy an animal uses must depend on its ability to collect accurate information on the risks to which it is exposed. However, models attempting to predict when a prey animal should escape from an approaching predator often assume that the prey has accurate information on a predators distance and direction of approach. To test whether such models could be applied to a prey animal with restricted sensory capabilities, I explored the predator avoidance behaviour of the fiddler crab Uca vomeris by approaching crabs with small dummies intended to mimic a hunting tern. The crabs responded strongly and reliably to the simple dummies. They were clearly sensitive to risk and responded more often and earlier, the further away they were from their refuge. The probability of response was most strongly influenced by how directly and therefore how closely the dummies approached the crabs and reached 100% for very direct approaches. Surprisingly, however, the crabs responded later when the dummies approached them more directly. I argue that this counterintuitive result reflects a lack of reliable information on the predators distance and movement relative to the crab and its refuge, because like many small animals, crabs cannot measure the distance to a predator. I conclude that general models attempting to predict when animals should respond to an approaching predator need to incorporate the information that prey animals have available at the time of response.

Distribution of photoreceptor types in the retina of a marsupial, the tammar wallaby (Macropus eugenii)

Mammalian retinae generally contain low numbers of short-wavelength-sensitive cones (S-cones) and higher numbers of middle- to long-wavelength-sensitive cones (M-cones). Some recent studies found topographic differences between the different photoreceptor types and in some instances between photoreceptors and ganglion cells. To investigate this question further, we constructed topographical maps of the different photoreceptors found in an Australian marsupial, the tammar wallaby. We used two polyclonal antibodies that have been shown to label S-cones (JH455) or M-cones (JH492) in a range of mammals. In the tammar wallaby, the antisera clearly distinguish two cone types. JH455 recognizes a small subset of cones (S-cones) with a density of less than 500 cells/mm2 in the ventral retina. Their density increases towards the dorsal retina to about 1600-2000 cells/mm2. JH492 recognizes all remaining cones (M-cones), but also faintly labels most cone cells recognized by JH455. The distribution of M-cones, unlike that of the S-cones, shows a clear horizontal streak of high cell density through the central retina, just like the ganglion cells. Unlike the ganglion cells, however, the M-cones do not peak in the temporal retina but show a very broad peak (12,000-18,000 cells/mm2) in the central or even slightly nasal retina. Based on our findings, the retina of the tammar can be divided into three distinct regions: firstly, the dorsal retina, which has a low ganglion and low cone cell density but a high percentage of S-cones (30%), is thought to provide good spectral sensitivity; secondly, the central horizontal band of retina, which has a high ganglion and high cone cell density and therefore provides good spatial resolution; and thirdly, the ventral retina, which has a low ganglion cell but high cone cell density with few S-cones (5%) and is therefore thought to have a high contrast sensitivity but low acuity.

The variable colours of the fiddler crab Uca vomeris and their relation to background and predation

SUMMARY Colour changes in fiddler crabs have long been noted, but a functional interpretation is still lacking. Here we report that neighbouring populations of Uca vomeris in Australia exhibit different degrees of carapace colours, which range from dull mottled to brilliant blue and white. We determined the spectral characteristics of the mud substratum and of the carapace colours of U. vomeris and found that the mottled colours of crabs are cryptic against this background, while display colours provide strong colour contrast for both birds and crabs, but luminance contrast only for a crab visual system. We tested whether crab populations may become cryptic under the influence of bird predation by counting birds overflying or feeding on differently coloured colonies. Colonies with cryptically coloured crabs indeed experience a much higher level of bird presence, compared to colourful colonies. We show in addition that colourful crab individuals subjected to dummy bird predation do change their body colouration over a matter of days. The crabs thus appear to modify their social signalling system depending on their assessment of predation risk.

The twilight zone: ambient light levels trigger activity in primitive ants

Many animals become active during twilight, a narrow time window where the properties of the visual environment are dramatically different from both day and night. Despite the fact that many animals including mammals, reptiles, birds and insects become active in this specific temporal niche, we do not know what cues trigger this activity. To identify the onset of specific temporal niches, animals could anticipate the timing of regular events or directly measure environmental variables. We show that the Australian bull ant, Myrmecia pyriformis, starts foraging only during evening twilight throughout the year. The onset occurs neither at a specific temperature nor at a specific time relative to sunset, but at a specific ambient light intensity. Foraging onset occurs later when light intensities at sunset are brighter than normal or earlier when light intensities at sunset are darker than normal. By modifying ambient light intensity experimentally, we provide clear evidence that ants indeed measure light levels and do not rely on an internal rhythm to begin foraging. We suggest that the reason for restricting the foraging onset to twilight and measuring light intensity to trigger activity is to optimize the trade-off between predation risk and ease of navigation.

Claw waving display changes with receiver distance in fiddler crabs, Uca perplexa

Effective communication is critically dependent on the successful transfer of information and, because environmental and social conditions can affect signal transmission, animals should be able to adjust their signals to optimize reliability. We show, apparently for the first time in a movement-based signal, that visual displays are adjusted with respect to the distance of signal receivers. Not only does this show the ability of the fiddler crab to judge distance, but this also shows that signalling is context dependent on surprisingly fine spatial and temporal scales. We elicited courtship behaviour in the crabs with tethered females and simultaneously recorded the displays of males from above and from crab-eye level. As females approached, males increased signal intensity by shortening display duration and altered signal form by reducing the lateral movement component of the waving signal. We suggest that males tune their waving display depending on receiver distance (a) to balance energetic costs with reproductive benefits, (b) to alter the information content of the signal and (c) to avoid signal misinterpretation. Such fine-scale context sensitivity is likely to be far more widespread in animal communication than hitherto recognized from similar signal modifications in auditory communication.

Visual gaze control during peering flight manoeuvres in honeybees

As animals travel through the environment, powerful reflexes help stabilize their gaze by actively maintaining head and eyes in a level orientation. Gaze stabilization reduces motion blur and prevents image rotations. It also assists in depth perception based on translational optic flow. Here we describe side-to-side flight manoeuvres in honeybees and investigate how the bees’ gaze is stabilized against rotations during these movements. We used high-speed video equipment to record flight paths and head movements in honeybees visiting a feeder. We show that during their approach, bees generate lateral movements with a median amplitude of about 20 mm. These movements occur with a frequency of up to 7 Hz and are generated by periodic roll movements of the thorax with amplitudes of up to ±60°. During such thorax roll oscillations, the head is held close to horizontal, thereby minimizing rotational optic flow. By having bees fly through an oscillating, patterned drum, we show that head stabilization is based mainly on visual motion cues. Bees exposed to a continuously rotating drum, however, hold their head fixed at an oblique angle. This result shows that although gaze stabilization is driven by visual motion cues, it is limited by other mechanisms, such as the dorsal light response or gravity reception.