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Tardigrade evolution Fossil Tardigrades • Tardigrades have a long evolutionary history • Specimens have been recovered from Siberia that are thought to be 530myo • This has given tardigrades time to develop a variety of forms as they have invaded a range of habitats (marine terrestrial). Cope’s Rule • Body size tends to increase in an animal lineage during its evolution. – eg. horses – eg dinosaurs Cope’s Rule - tardigrades Body length (µm) 400 300 200 100 A B C Assumes evolutionary sequence from A C and that examples drawn are representative of each group. No support is available from the fragmentary fossil record. Terrestrial invasion Terrestrial habitats are less regular and fluctuate more than marine habitats: • Seasonal changes • Diurnal changes • Unpredictable changes Resistant states • Anoxybiosis • Reaction to reduction in oxygen tension • Encystment • Reaction to slow changes in environment • Osmobiosis • Reaction to increased salinity • Cryobiosis • Reaction to freezing • Anhydrobiosis • Reaction to loss of water Anhydrobiosis • Reduction in body size – ‘tun’ formation • Invagination of extremities - legs • Folding of intersegmental cuticle to reduce exposed surface area • Loss of water – synthesis of trehalose Tun formation Intersegmetal folds Trehalose • • • • • Disaccharide Inert Stable Replaces water Reversible reaction Morphological adaptation Marine species exhibit a number of extravagant appendages that would be an impediment in a changeable terrestrial environment. During desiccation, they would be difficult fold up to reduce surface area and would be difficult to protect in the dry state – they could become brittle. The cephalic sensory organs would be the key organs – these are reduced or even internalised in terrestrial species. In summary Tardigrades have made the transition from marine habitats (where constant environments have supported diversity) to terrestrial habitats (where fluctuating environments have had an inhibiting effect upon Cope’s Rule as a result of the structural and physiological requirements for anhydrobiosis). The question remains: why have the ‘ancient’ marine groups not developed larger body size? References Kinchin, I.M. (1994) The Biology of Tardigrades. London, Portland Press. pp. 1 – 186. Kinchin, I.M. (1995) Evolutionary trends in the tardigrades. The Quekett Journal of Microscopy, 37(6): 493 – 498.