Adaptations of Crustaceans to Land: A Summary and Analysis of New Findings

Crustaceans have adapted to land through various morphological,physiological, biochemical, and behavioral modifications, ofwhich some are shared by all land-dwelling crustaceans and othersare unique to animals within a particular habitat. Among thethree groups of crustaceans having truly terrestrial members,the amphipods have achieved their success on land primarilyby behavioral means, while the isopods and the decapods havedeveloped many morphological, physiological, and biochemicaladaptations as well. In all three groups, behavioral modifications ensure that lossof water is minimal, that the animals are exposed to favorablerather than extreme environmental conditions, and that the fineline between evaporative cooling and excessive dehydration ismaintained. In most crustaceans the excretion of nitrogenous wastes requiresthat copious supplies of water be available for washing awaythe soluble end-products. Yet terrestrial isopods are able toexcrete ammonia as a gas, without being subject to toxic side-effects.In decapods, either ammonia or insoluble uric acid may be excreted,with ammonia the more likely product when water is available,uric acid when water is scarce. In adult land crabs water balance is maintained through theconcerted action of gills, pericardial sacs, and gut. Theseorgans may take up, store, and redistribute salts and waterin response to control exerted by the central nervous systemthrough its secretory products. In larvae of land crabs theseorgans are not known to function in this way. Rather, the larvaeare adapted to cope with osmotic problems of their planktonicexistence. Gaseous exchange in adult land crabs is carried on not onlyby the gills but also by the highly vascularized lining of thebranchial chambers, and the hemocyanin ot these crabs is adaptedto function in the environment peculiar to each species. Terrestrialcrabs seem unable to withstand low temperatures, but their highrate of cytochrome c oxidase activity may help them to survivewhen temperatures are high. Modifications in behavior must have occurred quite early inthe transition of crustaceans from sea to land. Then, as now,appropriate behavioral responses to light, temperatuie, humidity,tidal cycles, and so on. were crucial if a terrestrial animalwas to survive. Social interactions, both for courtship andfor aggression, required the sending and receiving of appropriatevisual and acoustic signals and were promoted by the ritualizationof potentially injurious patterns of behavior.