Todd R. Jackman and David B. Wake,
1994.
Evolutionary and Historical Analysis of Variation in the
Blotched Forms of Salamanders of the Ensatina Complex (Amphibia: Plethodontidae).
Evolution 48(3),1994,pp 876-897
ABSTRACT
Geographic variation in 23 to 29 protein-encoding genetic
loci was examined in 48 populations of the Ensatina complex, a "ring
species" distributed around the Central Valley of California. The samples
span two critical links in the chain of morphologically distinct units:
the transition from the unblotched to blotched color pattern types in the
vicinity of Lassen Peak, northeastern California, and a geographic gap in
the range of the complex in the San Gabriel Mountains, southern California.
A general pattern of isolation by distance with a regular buildup of genetic
distance correlated with increases in geographic distance characterizes
the populations studied, with the exception of a little-differentiated group
of populations in the northern Sierra Nevada; this region is postulated
to be a zone of genetic reticulation characterized by relatively high gene
flow. An adaptively significant color pattern is thought to have spread
into the northern Sierra Nevada from the south, but protein variants have
been introduced both from the north and the south. Genetic distances across
the San Gabriel Mountain gap match expectations from the pattern of buildup
of genetic distance as a function of geographic distance elsewhere in the
complex. A phylogenetic analysis of the protein data supports the reticulation
hypothesis; whereas the southernmost populations currently do constitute
a monophyletic assemblage, an "extinction experiment" demonstrates
that the distinction could be the result C':' the recent extinction of populations
in a present gap in our sampling. The Ensatina complex appears to be a dynamic
entity representing several stages in the evolution of species. It is a
ring species, and whereas various taxonomic arrangements are possible, no
taxonomic changes are proposed.
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