works for animals that are no more than a couple of millimeters thick, but for larger animals a respiration system was necessary. The rise of the arthropods has been an enduring problem for paleobiologists. Why was the arthropod so successful, particularly in the beginning? Segmented animals dominated Cambrian seas, and segmentation provides for repeated features. Segments obviously became important for locomotion but, for arthropods, segmentation appears to have conferred the more important advantage of distributed oxygen absorption. Each trilobite leg had an attached gill, and leg motion constantly drew fresh oxygenated water over each gill. Arthropods never developed the kinds of lungs that vertebrates have, or the pump gills of fish and other aquatic animals. Early arthropods breathed by moving their legs. Peter Ward’s recent hypothesis is that segments were first used for respiration, to provide a large gill surface area, and using the segments for locomotion came later. For trilobites, the same functionality that pushed water over gills was also coopted for food intake. Also, the leg-mounted gill was necessary because of an arthropod’s body armor; oxygen could not be absorbed through tough exoskeletons.


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