Locomotion in Anaspides (Anaspidacea, Malacostraca) - insights from a morpho-functional study of thoracopods with some observations on swimming and walking.

Anaspidacea is probably the most enigmatic higher taxon within the Malacostraca. Representatives of the genus Anaspides are often considered "living fossils" due to their strong resemblance to their Triassic relatives. In comparison to other extant Malacostraca, they possess a high number of plesiomorphic characters. Anaspidacea are often combined with another freshwater taxon, Bathynellacea, into the Syncarida, but actual evidence for monophyletic Syncarida is weak. Other potential sister groups are Eucarida, Euphausiacea, or Euphausiacea + Peracarida. Like Euphausiacea, Mysidacea and "natant" decapods, Anaspidacea show what has been called a "caridoid facies". In malacostracan crustaceans, a prominent mode of locomotion is swimming, as it almost certainly was for the malacostracans' common ancestor. However, when considered in detail, swimming is found to take quite different forms in the various taxa. To obtain a better understanding of the evolution of swimming, we analyzed locomotion in several species of Anaspides endemic to Tasmania. The morphology of the protopodal and exopodal musculature as well as the exo- and endoskeletal structures of the thoracopods were examined using 3D reconstruction of CLSM- and μCT data. Various aspects of locomotion were documented and described using recording techniques including in-habitat underwater filming and macro high-speed recording. Our analysis shows the high level of complexity of the muscular arrangement and skeletal construction in the thoracopods of these - in many regards plesiomorphic - malacostracans, and demonstrates the presence of epipodal musculature. Our recordings provide insight into various aspects of locomotion of Anaspides, including the congruence of locomotive kinematics in swimming and walking. Our morphological findings and observations on locomotion in Anaspides are compared to findings in other malacostracan shrimps and discussed in a phylogenetic context, with our analysis providing further support for Xenommacarida (sensuRichter, 1999: Anaspidacea + Euphausiacea + Peracarida). Further, a new hypothesis for the origin of the ventral brood pouch, the marsupium, of Peracarida is discussed. On the basis of a comparison of their morphology and biology, the oostegites which form the marsupium are suggested to be derived thoracopodal endites as present in female Anaspides. Finally, an evolutionary scenario is presented and mapped on a cladogram in which the evolution of malacostracan swimming from a common caridoid ancestor up to the different taxa present today is discussed.