Ensatina's basic story was laid out by Robert Stebbins 30 years before

Tom was born in 1977. Based on the ring-like distribution of the di"erent
forms, Robert had proposed that the species started o" in Northern
California and Oregon and then spread south along both sides of the
Central Valley, which was too dry and hot for salamanders.1
According to Robert's hypothesis, as the pioneering populations moved
south, they evolved into several subspecies with new color patterns and
adaptations for living in di"erent environments. By the time they met
again in Southern California as the subspecies eschscholtzii and klauberi,
he argued, they had each evolved so much that they no longer interbred
even though the subspecies blended into one another around the rest
of the ring. Since species are often dened by their inability to interbreed
with other species, Ensatina seemed to represent the whole process of
speciation all the gradual changes that accumulate in two lineages and
that wind up making them incompatible with one another.

Of course, since this all would have happened millions of years ago,
Robert wasn't around to observe any of it. He based his ideas on the
morphology, or body form, of the subspecies in this case, their color
patterns. First, neighboring subspecies were more similar to one another
than to those across the ring and seemed to blend into one another. From
this, he hypothesized that Ensatina represented a ring species. Robert
also noticed that the northern coastal form, called picta, had a pattern of
colors that seemed to encompass the other subspecies. It was easy to
imagine how the more specialized southern forms could have evolved
from picta. Based on this, Robert hypothesized that the two southward-
moving Ensatina lineages had both emerged from picta's immediate
ancestors.