Locomotion in Nebalia bipes: a possible model for Palaeozoic phyllocarid crustaceans

The morphofunctional aspects of locomotion in Recent phyllocarid crustaceans are presented, based on combined video observations of living specimens of the cosmopolitan nektobenthic Nebalia (Leptostraca, Nebaliidae) and scanning electron microscopy of fixed material. N. bipes is infaunal by day, showing a preference for organic-rich black muds and dim light conditions. Under natural conditions, emergence from sediment and nocturnal swimming activities are controlled by a circadian rhythm. The 1st and 2nd antennae perform the major role in digging. The carapace rostral plate acts as a ram diverting particles and preventing inputs of sediment into the carapace. Superficial burrowing may provide camouflaged shelter against predation. Swimming is accomplished by the combined action of the first four pairs of pleopods (backward metachronal flexion during the power stroke) and the well-articulated abdominal shaft (downward flap). Plumose setae present on both the pleopods and furcal rami (governed by low Reynolds numbers) behave like paddles maximizing the resistance to the water. Clusters of cuticular microscales (3-5 μm) and microdenticles are present on the external surface of the carapace and the trunk segments, respectively. Both show a uniform directional arrangement suggesting functional interpretations in relation to locomotion (e.g., to prevent back slippage during digging, to reduce turbulence in the flow layer close to the body, and/or to serve as mechanoreceptors for the detection of variation in water flow). Key features of functional importance in the locomotion of living leptostracans (natatory pleopods, a highly flexible abdomen, setulate or paddle-like furcal rami) are also recognized in Palaeozoic archaeostracan phyllocarids, suggesting that swimming was accomplished by the same pleopodal movements as described for N. bipes. Comparisons with Recent analogues (Nebaliopsis) suggest that phyllocarids with a free swimming life-style existed among the lower Palaeozoic archaeostracans (e.g., Caryocaris; Ordovician). Asymmetrical cuticular ornament in some archaeostracans indicates burrowing habits. Some Palaeozoic phyllocarids may have been occasional (possibly diurnal) mud dwellers comparable to modern nebaliids.