The Influence of Environmental and Physiological Factors on Variation in American Toad (Anaxyrus americanus) Dorsal Coloration
Adaptive coloration in amphibians is widespread and aids in camouflage, communication, and thermoregulation. Understanding the environmental factors that contribute to color variation is important for predicting how changes in climate and habitat parameters may alter amphibian fitness. Studies on toad coloration have focused on genetic elements, dorsal spot patterns, and juveniles, but less is known about the relationship between adult toad coloration and environmental conditions. The goal of this study was to examine dorsal color variation in adult American Toads (Anaxyrus americanus) in Erie County, Pennsylvania, USA, to determine how environmental and morphological factors influence coloration. We conducted visual surveys to sample adult toads across a habitat gradient and recorded several potential predictor variables (i.e., site elevation, substrate type, snout–vent length, and body surface temperature). We calibrated photographs of each toad and quantified red (R), green (G), and blue (B) color values within seven dorsal body regions. We summarized RGB values for each dorsal body region using a principal component analysis and used model selection approaches to select between models containing different predictor variables. The most supported model to explain the variation in color of all dorsal body regions contained only site elevation. On average, the body regions of all toads from higher elevation sites were darker than those of toads from lower elevations, but the amount of variation in dorsal coloration accounted for by elevation was low. Our results suggest that the factors that drive variation in toad dorsal coloration are complex, but that this trait is potentially sensitive to environmental changes.ABSTRACT

Examples of American Toad (Anaxyrus americanus) color variation found within the Wintergreen Gorge area of Erie, Pennsylvania, USA. Photographs (A)–(G) are of adult American Toads with dorsal pigmentation ranging from light tan to charcoal gray. Photo credits: M. Mack.

Dorsal regions sampled for color analysis and number of sample points measured per region: 1 = midline (four points); 2 = back spots (six points, three from each side of the midline); 3 = sides (three points per side); 4 = dark leg band (one point per hind leg); 5 = light leg band (one point per hind leg); 6 = leg spot (one point per wart/spot on each hind leg); 7 = parotid (one point per parotid gland). Photo credits: M. Mack.

American Toad (Anaxyrus americanus) dorsal coloration: a visualization of the range of color variation quantified for all sampled individuals. This plot contains one point per sampled dorsal body region of each toad and represents averaged R, G, and B values for that region. Lower numbers on all three axes represent darker colors, whereas higher values indicate lighter colors. The color gradient depicted approximates the range of colors present in all dorsal body regions of the toads used in this study.

Dorsal coloration by body region: Panels (A)–(G) show the color of each body region on R, G, and B axes for higher elevation toads (gray circles) and lower elevation toads (orange triangles), with each point representing an averaged R, G, and B value of each toad for the specified dorsal body region.
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