Abstract

Abstract To understand the function of colour signals in nature, we require robust quantitative analytical frameworks to enable us to estimate how animal and plant colour patterns appear against their natural background as viewed by ecologically relevant species. Due to the quantitative limitations of existing methods, colour and pattern are rarely analysed in conjunction with one another, despite a large body of literature and decades of research on the importance of spatio‐chromatic colour pattern analyses. Furthermore, key physiological limitations of animal visual systems such as spatial acuity, spectral sensitivities, photoreceptor abundances and receptor noise levels are rarely considered together in colour pattern analyses. Here, we present a novel analytical framework, called the Quantitative Colour Pattern Analysis (QCPA). We have overcome many quantitative and qualitative limitations of existing colour pattern analyses by combining calibrated digital photography and visual modelling. We have integrated and updated existing spatio‐chromatic colour pattern analyses, including adjacency, visual contrast and boundary strength analysis, to be implemented using calibrated digital photography through the Multispectral Image Analysis and Calibration (MICA) Toolbox. This combination of calibrated photography and spatio‐chromatic colour pattern analyses is enabled by the inclusion of psychophysical colour and luminance discrimination thresholds for image segmentation, which we call ‘Receptor Noise Limited Clustering’, used here for the first time. Furthermore, QCPA provides a novel psycho‐physiological approach to the modelling of spatial acuity using convolution in the spatial or frequency domains, followed by ‘Receptor Noise Limited Ranked Filtering’ to eliminate intermediate edge artefacts and recover sharp boundaries following smoothing. We also present a new type of colour pattern analysis, the ‘local edge intensity analysis’ as well as a range of novel psycho‐physiological approaches to the visualization of spatio‐chromatic data. QCPA combines novel and existing pattern analysis frameworks into what we hope is a unified, free and open source toolbox and introduces a range of novel analytical and data‐visualization approaches. These analyses and tools have been seamlessly integrated into the MICA toolbox providing a dynamic and user‐friendly workflow.

Highlights

  • Animal colour patterns are complex traits which serve a multitude of purposes, including defence against predators, social signalling and thermoregulation (Cott, 1940)

  • We present a new type of colour pattern analysis, the

  • We introduce a method to overcome these problems and present a user-friendly, open-source framework, which we call ‘Quantitative Colour Pattern Analysis’ (QCPA)

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Summary

Introduction

Animal colour patterns are complex traits which serve a multitude of purposes, including defence against predators (such as camouflage and aposematism), social signalling and thermoregulation (Cott, 1940). Spatiochromatic colour pattern analyses overcome these limitations as they are designed to consider perceptual interactions between spatial, chromatic and achromatic information (Endler, 1978, 1990; Vorobyev et al, 2001; Endler & Mielke, 2005; Marshall et al, 2006; Stevens & Merilaita, 2011; Endler & Mappes, 2017; Olsson et al, 2017; Endler et al, 2018; Ruxton et al, 2018) Such approaches parameterise the properties of colour patterns, including colour adjacency, pattern regularity, visual contrast and colour pattern similarity Potential applications of QCPA include (but are not limited to): background matching, disruptive colouration, polymorphism, mimicry, aposematism, sexual signalling, territorial signalling, thermoregulation and landscape analysis

Materials and Methods
6.1: Adjacency Analysis
6.2: Visual Contrast Analysis
6.3: Boundary Strength Analysis
Discussion
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