On Papuliferes, putative ciliate cysts of diverse morphologies, with new observations from the plankton of the Chukchi Sea (Arctic Ocean)

In the early 20th century, Alphonse Meunier described "Papuliferes" as a group of enigmatic forms of unknown taxonomic affinity characterized by possessing a hyaline pimple, a "papula". In the early 1980's Papuliferes were equated with cysts of tintinnid ciliates. The most conspicuous forms, the large Fusopsis, have been widely found, and are now known to resemble the cyst of a certain species of oligotrich ciliate (i.e., Cyrtostrombidium boreale). Thus today, Papulifere forms are often assumed to be cysts of oligotrich ciliates. Here we report on 26 Papulifere forms, of more or less distinct morphologies, found in the plankton of the Chukchi Sea. We found forms resembling some of those described by Meunier, and recorded here for the first time since Meunier's reports, and others that do not resemble any of Meunier's Papuliferes. Here, we first review the literature on Papuliferes, then we present the surprising variety of forms we found in Chukchi Sea, and for some, we provide for the first time data on morphological variability. With this report we have expanded the catalogue of observed Papulifere forms and documented variability in the dimensions of some morphotypes. However, we urge caution in assigning a ciliate species name to any given Papulifere form in the absence of corroborating data. There is a need for observational and/or sequencebased data to elucidate the identity of Papulifere forms.