Nematodes are widely thought to be 'eutelic', that is, each adult tissue is made up of an invariant number of cells. We wondered whether this was always true, and so counted the numbers of cells in the hypodermal syncytium (
hyp-7) in 15 different free-living species (Rhabditids, Panagrolaimids, and Cephalobids). Among species, mean hypodermal cell number varies 3-fold, but cellular constancy (as measured by among-individual variance in cell number) varies by 3 orders of magnitude. Furthermore, there is a strong positive correlation between the mean and variance in cell number. Thus, Panagrellus redivivus has approximately 3x as many hypodermal cells as C. elegans , but is 300x as variable. We reasoned that there are two basic ways in which one species might come to be more variable than another. First, it could be some species are more variable just because they have more complex lineages. Second, it could be that some species are more variable because they have a higher error rate during particular cellular events. To test among these alternatives, we first determined the canonical V cell lineages of two species, Oscheius myriophila and Rhabditella octopleura . Then we modelled these lineages (and those of C. elegans , and P. redivivus ) computationally, introducing plausible kinds of variation in cell divisions and fates into the simulation. These simulations generate predictions, for a given canonical lineage, of the relationship between cellular error rates and variance in adult cell number. We found that the enormous variance observed in P. redivivus (and in some other species such as Rhabditoides regina ) can only be explained by far higher error rates in V cell lineages than are found in C. elegans . Notably, Sternberg and Horvitz, when lineaging P. redivivus , observed variant V cell lineages of a sort that have never been found in C. elegans .