Adaptive divergence and speciation may happen despite opposition by gene flow. Identifying the genomic basis underlying divergence with gene flow is a major task in evolutionary genomics. Most approaches (e.g. outlier scans) focus on genomic regions of high differentiation. However, not all genomic architectures potentially underlying divergence are expected to show extreme differentiation. Here, we develop an approach that combines hybrid zone analysis (i.e. focuses on spatial patterns of allele frequency change) with system-specific simulations to identify loci inconsistent with neutral evolution. We apply this to a genome-wide SNP set from an ideally-suited study organism, the intertidal snail Littorina saxatilis, which shows primary divergence between ecotypes associated with different shore habitats. We detect many SNPs with clinal patterns, most of which are consistent with neutrality. Among non-neutral SNPs, most are located within three large putative inversions differentiating ecotypes. Many non-neutral SNPs show relatively low levels of differentiation. We discuss potential reasons for this pattern, including loose linkage to selected variants, polygenic adaptation and a component of balancing selection within populations (which may be expected for inversions). Our work is in line with theory predicting a role for inversions in divergence, and emphasises that genomic regions contributing to divergence may not always be accessible with methods purely based on allele frequency differences. These conclusions call for approaches that take spatial patterns of allele frequency change into account in other systems.
Cline fitting results
Provides information about all SNPs (one SNP per line) that passed filters and where either a model of clinal change provided the best fit (Type=Cline), or the allele frequency difference between cline ends was < 0.1 / the model of no clinal change provided the best fit (Type=NC).
- Contig: Reference genome contig the SNP is located on
- Position: Position of the SNP within the contig
- Type: Whether SNP showed clinal change (Cline) or no change (NC)
- Wave: Whether the reference allele is the more common allele in the Wave ecotype (1) or not (2)
- Centre: Position of the cline centre on the path along the shore (in m)
- Width: Width of the cline (in m)
- p_crab: Allele frequency in the Crab ecotype
- p_diff: Difference between Crab and Wave allele frequency
- Var.Ex: Variance explained by the cline model
- AIC_NC, AIC_cline, AIC_lt, AIC_rt, AIC_bt: AICs for different models (NC, Cline, left-tailed cline, right-tailed cline, two-tailed cline)
- sel: Whether a cline is inconsistent with neutrality, indicating direct or linked selection (i.e. whether Var.Ex is > then the threshold [35.69] defined by simulations)
- LG: Linkage group from genetic map on which SNP is located
- av: Position on the linkage group where SNP is located
WestramEtAl2018_cline_results_threshold35.69.txt
Capture probe sequences
List of all 40,000 capture sequencing probes used in this study. Note that the probe IDs refer to previous L. saxatilis assemblies and are not meaningful with respect to position in the current assembly.
WestramEtAl2018_probe_sequences.txt
Snail positions and phenotypes
Provides information about the sampling coordinates and phenotypes of each snail.
- x, y, z: Position of snail on the shore in 3 dimensions
- RW1, RW2: Relative warps of geometric morphometric analysis (shell shape)
- length_mm: Shell length (in mm)
- CS: Centroid size from geometric morphometric analysis
- DistAlongPath_m: Position on the path along the shore (in m)
WestramEtAl2018_snail_positions_phenotypes.csv
Littorina saxatilis genome assembly
L. saxatilis genome assembly used in this study (repeat regions hard-masked).
WestramEtAl2018_Littorina_saxatilis_assembly_hard_masked.zip