title: Strong selection against hybrids at a hybrid zone in the Ensatina ring species complex and its evolutionary implications.
abstract: The analysis of interactions between lineages at varying levels of genetic divergence can provide insights into the process of speciation through the accumulation of incompatible mutations. Ring species, and especially the Ensatina eschscholtzii system exemplify this approach. The plethodontid salamanders E. eschscholtzii xanthoptica and E. eschscholtzii platensis hybridize in the central Sierran foothills of California. We compared the genetic structure across two transects (southern and northern Calaveras Co.), one of which was resampled over 20 years, and examined diagnostic molecular markers (eight allozyme loci and mitochondrial DNA) and a diagnostic quantitative trait (color pattern). Key results across all studies were: (1) cline centers for all markers were coincident and the zones were narrow, with width estimates of 730 m to 2000 m; (2) cline centers at the northern Calaveras transect were coincident between 1981 and 2001, demonstrating repeatability over five generations; (3) there were very few if any putative F1s, but a relatively high number of backcrossed individuals in the central portion of transects; and (4) we found substantial linkage disequilibrium in all three studies and strong heterozygote deficit both in northern Calaveras, in 2001, and southern Calaveras. Both linkage disequilibrium and heterozygote deficit showed maximum values near the center of the zones. Using estimates of cline width and dispersal, we infer strong selection against hybrids. This is sufficient to promote accumulation of differences at loci that are neutral or under divergent selection, but would still allow for introgression of adaptive alleles. The evidence for strong but incomplete isolation across this centrally located contact is consistent with theory suggesting a gradual increase in postzygotic incompatibility between allopatric populations subject to divergent selection and reinforces the value of Ensatina as a system for the study of divergence and speciation at multiple stages.
authors.affiliation: Museum of Vertebrate Zoology, University of California, Berkeley, 3101 Valley Life Science Building 3160, Berkeley, California 94720-3160, USA. jalex@rc.unesp.br
authors.forename: João
authors.initials: J
authors.lastname: Alexandrino
authors.forename: Stuart J E
authors.initials: SJ
authors.lastname: Baird
authors.forename: Lucinda
authors.initials: L
authors.lastname: Lawson
authors.forename: J Robert
authors.initials: JR
authors.lastname: Macey
authors.forename: Craig
authors.initials: C
authors.lastname: Moritz
authors.forename: David B
authors.initials: DB
authors.lastname: Wake
mesh_terms: D000818:Animals; D005075:Biological Evolution; D002140:California; D045303:Cytochromes b / Q000235:genetics; D006580:Genetic Carrier Screening; D005828:Genetics, Population; D005843:Geography; D006824:Hybridization, Genetic; D007527:Isoenzymes; D016013:Likelihood Functions; D015810:Linkage Disequilibrium; D008137:Longitudinal Studies; D010858:Pigmentation / Q000502:physiology; D012150:Polymorphism, Restriction Fragment Length; D012641:Selection, Genetic; D013045:Species Specificity; D014562:Caudata / Q000235:genetics* / Q000502:physiology
chemical_list: D007527:Isoenzymes; D045303:Cytochromes b