Phylogeny and evolution of apusomonadida (protozoa: apusozoa): new genera and species.

Apusomonadida (Apusomonas; Amastigomonas) are understudied gliding zooflagellates. We divide Amastigomonas into five genera, three new: Podomonas; Manchomonas; Multimonas. Microscopy and 18S rDNA sequences establish three new marine species (Podomonas magna; P. capensis; Multimonas media) and a new cyst-forming non-marine species from the surface of ivy leaves (Thecamonas oxoniensis). We consider the soil and freshwater Amastigomonas debruynei, caudata, and borokensis generically distinct from marine Thecamonas. We establish the new combination Multimonas marina (formerly Cercomonas or Amastigomonas). We studied by DIC microscopy and 18S rDNA sequencing three strains microscopically indistinguishable from marine Thecamonas trahens and argue that marine strains of almost identical sequence and appearance (visible largely acronematic cilia) were previously misidentified as Am. debruynei. We argue that 'Amastigomonas sp.' ATCC50062, whose 18S rRNA was sequenced previously and whose complete genome is being sequenced, is T. trahens. We include electron micrographs of T. aff. trahens, P. capensis and magna; ultrastructural cytoskeletal differences between P. capensis, Thecamonas, and Manchomonas (=Amastigomonas) bermudensis comb. n. allow novel functional interpretations of apusomonad evolution. On 18S rDNA trees Apusomonas and Manchomonas form a robust clade (Apusomonadinae), but Thecomonas trahens, T. oxoniensis, Multimonas, and Podomonas all branch deeply but unstably. Apusomonadida and Planomonas are weakly sister to opisthokonts.