To be effective, animal colour signals must attract attention—and therefore need to be conspicuous. To understand the signal function, it is useful to evaluate their conspicuousness to relevant viewers under various environmental conditions, including when visual scenes are cluttered by objects of varying colour. A widely used metric of colour difference (Δ S ) is based on the receptor noise limited (RNL) model, which was originally proposed to determine when two similar colours appear different from one another, termed the discrimination threshold (or just noticeable difference). Estimates of the perceptual distances between colours that exceed this threshold—termed ‘suprathreshold’ colour differences—often assume that a colour's conspicuousness scales linearly with colour distance, and that this scale is independent of the direction in colour space. Currently, there is little behavioural evidence to support these assumptions. This study evaluated the relationship between Δ S and conspicuousness in suprathreshold colours using an Ishihara-style test with a coral reef fish, Rhinecanthus aculeatus . As our measure of conspicuousness, we tested whether fish, when presented with two colourful targets, preferred to peck at the one with a greater Δ S from the average distractor colour. We found the relationship between Δ S and conspicuousness followed a sigmoidal function, with high Δ S colours perceived as equally conspicuous. We found that the relationship between Δ S and conspicuousness varied across colour space (i.e. for different hues). The sigmoidal detectability curve was little affected by colour variation in the background or when colour distance was calculated using a model that does not incorporate receptor noise. These results suggest that the RNL model may provide accurate estimates for perceptual distance for small suprathreshold distance colours, even in complex viewing environments, but must be used with caution with perceptual distances exceeding 10 Δ S .
In cooperatively breeding species, subordinates can obtain group membership through social interactions with other group members or by providing services such as helping with territory defence. Large subordinate individuals, which can reproduce, are expected to adjust their behaviour as a function of the demand of help and group size because if the environmental conditions allow, they may either leave the group to start breeding or queue for the breeding position in their natal group. The number of helpers in a group is expected to affect the need of help by dominants and consequently also the level of subordination shown by helpers. In a series of field experiments, we manipulated the need of help and the opportunities for subordinates to show submissive behaviour in a wild population of the cooperatively breeding species Neolamprologus pulcher. We assessed if group size determines the social behavioural strategy of large subordinate individuals. When experimentally eliciting submissive behaviour, large subordinates from small groups showed a lower frequency of submissive behaviour compared to large groups; moreover, they tended to show a higher frequency of sand digging than in large groups. In contrast, neither territory defence in the presence of a heterospecific egg and larvae predator nor dispersal propensity, measured as prospecting frequency in neighbouring territories, was affected by group size. A principal component analysis revealed that prospecting is uncorrelated with submissive behaviour and helping behaviour. Our results suggest that group size may be involved in shaping behavioural phenotypes of juvenile subordinates.
Predators learn and memorise the association between conspicuous colour patterns of aposematic prey and their underlying chemical or secondary defences. Consequently, variation in signal design within a species should be selected against, because it can decrease the rate of predator learning and enhance the rates of predator errors. However, quantitative assessments have not been made on whether the strength of chemical defences influences colour pattern variation. We examined this by quantifying colour pattern variation using Quantitative Colour Pattern Analysis (QCPA) in 12 Dorid nudibranch species (Infraorder: Doridoidei) that varied in their unpalatability. We accounted for the physiological limitations of a potential predator visual system (a triggerfish,Rhinecanthus aculeatus), modelling animal appearance along an escalating predation sequence. We found that various colour pattern statistics were less variable in highly unpalatable species, with pattern statistics being up to 72% less variable than those of more palatable species. No correlations indicating the opposite were found across 157 colour pattern statistics. However, the strength and number of correlations depended on viewing distance. Our results suggest that low colour pattern variability could be favourable for aposematic signalling in Dorid nudibranchs. We provide evidence for distance-dependent signalling facilitating context-specific feature selection in multicomponent visual signals.
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