There is a bewildering diversity of signals, sensory systems, and signaling behavior. A consideration of how these traits affect each other's evolution explains some of this diversity. Natural selection favors signals, receptors, and signaling behavior that maximize the received signals relative to background noise and minimize signal degradation. Properties of sensory systems bias the direction of evolution of the signals that they receive. For example, females may prefer males whose signals they can perceive more easily, and this will lead to the spread of more easily perceived male traits. Environmental conditions during signal transmission and detection also affect signal perception. Specific environmental conditions will bias the evolutionary direction of behavior, which affects the time and place of signaling as well as microhabitat preferences. Increased specialization of microhabitats and signaling behavior may lead to biased evolution of the sensory systems to work more efficiently. Thus, sensory systems, signals, signaling behavior, and habitat choice are evolutionarily coupled. These suites of traits should coevolve in predictable directions, determined by environmental biophysics, neurobiology, and the genetics of the suites of traits-hence the term "sensory drive." Because conditions vary in space and time, diversity will be generated.

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In studies of animal colouration i t is no longer necessary to rely on subjective assessments of colour and conspicuousness, nor on methods which rely upon human vision. This is important because animals vary greatly in colour vision and colour is context-dependent. New methods make i t practical to measure the colour spectrum of pattern elements (patches) of animals and their visual backgrounds for the conditions under which patch spectra reach the conspecific's, predator's or prey's eyes. These methods can be used in both terrestrial and aquatic habitats. A patch's colour is dependent not only upon its reflectance spectrum, but also upon the ambient light spectrum, the transmission properties of air or water, and the veiling light sprctrum. These factors change with time of day, weather, season and microhabitat, so colours must be measured under the conditions prevalent when colour patterns are normally used. Methods of measuring, classifying and comparing colours are presented, as well as techniques for assessing the conspicuousness of colour patterns as a whole. Some implications of the effect of environmental light and vision are also discussed.

Forests exhibit much variation in light environments, and this can affect communication among animals, communication between animals and plants, photosynthesis, and plant morphogenesis. Light environments are caused by, and can be predicted from, the geometry of the light paths, the weather conditions, and the time of day. The structure of forests leads to four major light habitats when the sun is not blocked by clouds: forest shade, woodland shade, small gaps, and large gaps. These are characterized by yellow-green, blue-gray, reddish, and "white" ambient light spectra, respectively. When the sun is blocked by clouds, the spectra of these four habitats converge on that of large gaps and open areas, so the single light environment during cloudy weather will be called open/cloudy. An additional light environment (early/late) is associated with low sun angles (near dawn or dusk); it is purplish. Each light environment is well defined and was found in forests of Trinidad, Panama, Costa Rica, Australia, California, and Florida. Scattered literature references suggest similar patterns elsewhere in North America, Europe, and Java. Perceived colors of animals, flowers, and fruits depend upon the interaction between ambient light color and the reflectance color of the animal or plant parts. As a result, an animal or plant may have a different appearance in each environment, i.e., a color pattern may be relatively cryptic in some light environments while relatively conspicuous in others. This has strong implications for the joint evolution of visual signals and vision, as well as microhabitat choice. Plant growth and form may also be affected by variation in the color of forest light.

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Colour patterns and their visual backgrounds consist of a mosaic of patches that vary in colour, brightness, size, shape and position. Most studies of crypsis, aposematism, sexual selection, or other forms of signalling concentrate on one or two patch classes (colours), either ignoring the rest of the colour pattern, or analysing the patches separately. We summarize methods of comparing colour patterns making use of known properties of bird eyes. The methods are easily modifiable for other animal visual systems. We present a new statistical method to compare entire colour patterns rather than comparing multiple pairs of patches. Unlike previous methods, the new method detects differences in the relationships among the colours, not just differences in colours. We present tests of the method's ability to detect a variety of kinds of differences between natural colour patterns and provide suggestions for analysis. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86, 405–431.

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