The branching morphology of Ediacaran rangeomorph fronds has no exact counterpart in other complex macroorganisms. As such, these fossils pose major questions as to growth patterns, functional morphology, modes of feeding, and adaptive optimality. Here, using parametric Lindenmayer systems, a formal model of rangeomorph morphologies reveals a fractal body plan characterized by self-similar, axial, apical, alternate branching. Consequent morphological reconstruction for 11 taxa demonstrates an adaptive radiation based on 3D space-filling strategies. The fractal body plan of rangeomorphs is shown to maximize surface area, consistent with diffusive nutrient uptake from the water column (osmotrophy). The enigmas of rangeomorph morphology, evolution, and extinction are resolved by the realization that they were adaptively optimized for unique ecological and geochemical conditions in the late Proterozoic. Changes in ocean conditions associated with the Cambrian explosion sealed their fate.paleobiology | paleontology I n parallel with large-scale geochemical transitions associated with ocean oxygenation (1-3), the Ediacaran Period (635-541 Ma) records a major diversification of multicellular eukaryotes. Rangeomorph fronds (575-541 Ma) dominated early Ediacaran biotas (4) and have a characteristic branching morphology, distinct from any known Phanerozoic organism (5). Although the fronds are often preserved as flattened impressions, exceptional moldic fossils preserve details of the 3D branching structure to a resolution of 30 μm (6). Qualitative classifications for rangeomorph branching patterns have been proposed (7,8), but no quantitative model has previously been formulated. Because branching is repeated over decreasing size scales (with up to four observed orders of branching), rangeomorph fronds have been informally described as self-similar and fractal (4-6). Although this has potential implications for the functional optimality of their morphologies (5, 9), the extent to which they are formally fractal and self-similar (10, 11) has not previously been tested. Furthermore, until now, evolutionary transitions in branching patterns have not been characterized within any quantitative framework.Rangeomorphs inhabited shallow to abyssal marine environments (1,8,(12)(13)(14), evidently precluding photosynthesis for most taxa (12). Preservational features, including bending and overfolding (4, 15), suggest that rangeomorphs were soft-bodied. No evidence exists for either motility or active feeding (such as musculature, filter feeding organs, or a mouth). Consequently, rangeomorphs have been reconstructed as sessile, feeding on organic carbon by diffusion (or possibly endocytosis) through the body surface (3,5,12,16), with a large surface area to volume ratio aiding nutrient uptake (5, 16). The adaptive potential of their different branching morphologies has, however, never been quantified.Here, using parametric Lindenmayer systems (L-systems) (17, 18), we present a unified model to describe the branching structure of Ed...