約瑟夫·萊迪
The first person to observe and formally describe symbiotic flagellates in "R. flavipes" was the American paleontologist Joseph Leidy. In 1877, he reported his findings to the Academy of Natural Sciences of Philadelphia, describing three new species which he mistakenly took to be parasitic ciliates: Trichonympha agilis, Pyrsonympha vertens, and Dinenympha gracilis.