| 1. |
- Rolan-Alvarez, E., et al.
(author)
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Nonallopatric and parallel origin of local reproductive barriers between two snail ecotypes
- 2004
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In: Molecular Ecology. - 0962-1083. ; 13:11, s. 3415-3424
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Journal article (peer-reviewed)abstract
- Theory suggests that speciation is possible without physical isolation of populations (hereafter, nonallopatric speciation), but recent nonallopatric models need the support of irrefutable empirical examples. We collected snails (Littorina saxatilis) from three areas on the NW coast of Spain to investigate the population genetic structure of two ecotypes. Earlier studies suggest that these ecotypes may represent incipient species: a large, thick-shelled 'RB' ecotype living among the barnacles in the upper intertidal zone and a small, thin-shelled 'SU' ecotype living among the mussels in the lower intertidal zone only 10-30 m away. The two ecotypes overlap and hybridize in a midshore zone only 1-3 m wide. Three different types of molecular markers [allozymes, mitochondrial DNA (mtDNA) and microsatellites] consistently indicated partial reproductive isolation between the RB and the SU ecotypes at a particular site. However, each ecotype was related more closely to the other ecotype from the same site than to the same ecotype from another site further along the Galician coast (25-77 km away). These findings supported earlier results based solely on allozyme variation and we could now reject the possibility that selection produced these patterns. The patterns of genetic variation supported a nonallopatric model in which the ecotypes are formed independently at each site by parallel evolution and where the reproductive barriers are a byproduct of divergent selection for body size. We argue that neither our laboratory hybridization experiments nor our molecular data are compatible with a model based on allopatric ecotype formation, secondary overlap and introgression.
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| 2. |
- Butlin, Roger, 1955, et al.
(author)
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PARALLEL EVOLUTION OF LOCAL ADAPTATION AND REPRODUCTIVE ISOLATION IN THE FACE OF GENE FLOW
- 2014
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In: Evolution. - : Wiley. - 0014-3820. ; 68:4, s. 935-949
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Journal article (peer-reviewed)abstract
- Parallel evolution of similar phenotypes provides strong evidence for the operation of natural selection. Where these phenotypes contribute to reproductive isolation, they further support a role for divergent, habitat-associated selection in speciation. However, the observation of pairs of divergent ecotypes currently occupying contrasting habitats in distinct geographical regions is not sufficient to infer parallel origins. Here we show striking parallel phenotypic divergence between populations of the rocky-shore gastropod, Littorina saxatilis, occupying contrasting habitats exposed to either wave action or crab predation. This divergence is associated with barriers to gene exchange but, nevertheless, genetic variation is more strongly structured by geography than by ecotype. Using approximate Bayesian analysis of sequence data and amplified fragment length polymorphism markers, we show that the ecotypes are likely to have arisen in the face of continuous gene flow and that the demographic separation of ecotypes has occurred in parallel at both regional and local scales. Parameter estimates suggest a long delay between colonization of a locality and ecotype formation, perhaps because the postglacial spread of crab populations was slower than the spread of snails. Adaptive differentiation may not be fully genetically independent despite being demographically parallel. These results provide new insight into a major model of ecologically driven speciation.
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| 3. |
- Conde-Padin, P., et al.
(author)
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Revealing the mechanisms of sexual isolation in a case of sympatric and parallel ecological divergence
- 2008
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In: Biological Journal of the Linnean Society. - : Oxford University Press (OUP). - 0024-4066 .- 1095-8312. ; 94:3, s. 513-526
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Journal article (peer-reviewed)abstract
- Two ecotypes of a marine intertidal snail (Littorina saxatilis), living at different microhabitats and shore levels, have evolved in sympatry and in parallel across the Galician rocky shore. These ecotypes differ in many traits (including size) due to differential adaptation. They meet, mate assortatively, and partially hybridize at the mid shore where the two microhabitats overlap. The partial sexual isolation observed is claimed to be a side-effect of the size differences between ecotypes combined with a size assortative mating found in most populations of this species. We investigated this hypothesis using three complementary experimental approaches. First, we investigated which of the different shell variables contributed most to the variation in individual sexual isolation in the field by using two new statistics developed for that purpose: (1) pair sexual isolation and (2) r(i), which is based on the Pearson correlation coefficient. We found that size is the most important trait explaining the sexual isolation and, in particular, the males appear to be the key sex contributing to sexual isolation. Second, we compared the size assortative mating between regions: exposed rocky shore populations from north-westwern Spain (showing incomplete reproductive isolation due to size assortative mating) and protected Spanish and Swedish populations (showing size assortative mating but not reproductive isolation between ecomorphs). Most of the variation in size assortative mating between localities was significantly explained by the within-population level of variation on size. Third, we performed a laboratory male choice experiment, which further suggested that the choice is made predominantly on the basis of size. These results confirm the mechanism proposed to explain the sexual isolation in the Galician hybrid zone and thus support this case as a putative example of parallel incipient speciation. (C) 2008 The Linnean Society of London.
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| 4. |
- Erlandsson, Johan, et al.
(author)
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Migratory differences between ecotypes of the snail Littorina saxatilis on Galician rocky shores
- 1998
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In: Evolutionary Ecology. - 0269-7653. ; 12:8, s. 913-924
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Journal article (peer-reviewed)abstract
- Local migration patterns may be crucial to gene flow in species of marine gastropods which do not broadcast pelagic larvae. In some species, dispersal over distances of a few metres may influence micro-scale population structures. We investigated the migration pattern in Galician populations of the snail Littorina saxatilis in which populations of contrasting morphologies occupy different tidal levels of the same rocky shore. Two distinct morphs, one at the upper and one at the lower shore, overlap in distribution in a small mid-shore region where hybrids are produced. We documented the distances and directions of migration of both parental morphs and hybrids 1 month after they had been marked and released at different shore levels. When placed at their native shore level, snails migrated less than about 2 m and usually in independent directions. This supports the suggestion of a low local gene flow. At an alien shore level, however, the morphs often moved further and more directionally compared with native morphs. These differences may help to keep the two morphs separated at different shore levels. As fitness of an individual is highest in its native habitat, this seems to be an adaptive strategy.
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| 5. |
- Johannesson, Kerstin, 1955, et al.
(author)
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Hybrid fitness seems not to be an explanation for the partial reproductive isolation between ecotypes of Galician Littorina saxatilis
- 2000
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In: Journal of Molluscan Studies. - 0260-1230. ; 66, s. 149-156
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Journal article (peer-reviewed)abstract
- Partial reproductive isolation between closely related groups of organisms is suggested to be of central importance during speciation. Galician populations of Littorina saxatilis are phenotypically differentiated into an upper-shore and a lower-shore morph. These mate assortatively in the mid-shore: zone of overlap, and genetic assessment shows an impeded gene flow between the parental morphs. The traditional explanation as to why assortative mating occurs is that reproductive isolation is reinforced due to hybrid unfitness. Earlier studies have, however, not found hybrids to be less viable. Likewise, growth and migratory behaviours are merely intermediate between those of the parental morphs. In the present study we compared male and female fertility components of the parental morphs and the hybrids to test hypotheses of decreased hybrid fertility. The results showed that hybrid males were as fertile as other males, and hybrid females did not produce fewer embryos, nor aborted embryos at a higher rate, than the parental morphs.
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| 6. |
- Rivas, M. J., et al.
(author)
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Population genomics of parallel evolution in gene expression and gene sequence during ecological adaptation
- 2018
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In: Scientific Reports. - : Springer Science and Business Media LLC. - 2045-2322. ; 8:1
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Journal article (peer-reviewed)abstract
- Natural selection often produces parallel phenotypic changes in response to a similar adaptive challenge. However, the extent to which parallel gene expression differences and genomic divergence underlie parallel phenotypic traits and whether they are decoupled or not remains largely unexplored. We performed a population genomic study of parallel ecological adaptation among replicate ecotype pairs of the rough periwinkle (Littorina saxatilis) at a regional geographical scale (NW Spain). We show that genomic changes underlying parallel phenotypic divergence followed a complex pattern of both repeatable differences and of differences unique to specific ecotype pairs, in which parallel changes in expression or sequence are restricted to a limited set of genes. Yet, the majority of divergent genes were divergent either for gene expression or coding sequence, but not for both simultaneously. Overall, our findings suggest that divergent selection significantly contributed to the process of parallel molecular differentiation among ecotype pairs, and that changes in expression and gene sequence underlying phenotypic divergence could, at least to a certain extent, be considered decoupled processes. © 2018, The Author(s).
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| 7. |
- Rolan-Alvarez, E., et al.
(author)
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Mechanisms of incomplete prezygotic reproductive isolation in an intertidal snail: testing behavioural models in wild populations
- 1999
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In: J. Evol. Biol.. ; 12, s. 879-890
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Journal article (peer-reviewed)abstract
- Two morphs (ecotypes) of the marine snail Littorina saxatilis coexist along Galician exposed rocky shores. They hybridize, but gene flow is impeded by a partial prezygotic reproductive barrier, and we have earlier suggested that this is a case of incipient sympatric speciation. To assess the mechanisms of prezygotic reproductive isolation, we estimated deviations from random mating (sexual selection and sexual isolation) of sympatric snails in 13 localities on the shore, and performed mate choice experiments in the laboratory. We also investigated the microdistribution of both morphs over patches of barnacles and blue mussels in the hybridization zone. We used computer simulations to separate the mechanisms contributing to reproductive isolation. On the shores sampled, male-female pairs were strongly assortative both with respect to morphs (mean Yule's V = 0.77) and size (mean Pearson's r = 0.47). In the laboratory, males of both morphs mounted other snails and mated other males and juveniles at random. However, mature females of equal sizes mated assortatively with respect to morph. The two morphs were non-randomly distributed over barnacle and mussel patches in the hybridization zone. Monte Carlo simulations showed that this microdistribution could explain about half the morph and size relationships in male-female pairs, while a simple rejection mechanism, rejecting the first 1-3 mates if they were of contrasting morphs, accounted for the remaining part of the reproductive isolation, and for parts of the size relationships found between mates. A size discriminant mate choice mechanism may also, to a lesser extent, contribute to the sexual isolation. Sexual selection was observed for female size (larger ones being favoured) and among certain morphs, but distinct biological mechanisms may cause these processes.
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| 8. |
- Rolan-Alvarez, E., et al.
(author)
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The maintenance of a cline in the marine snail Littorina saxatilis: The role of home site advantage and hybrid fitness
- 1997
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In: Evolution. - 0014-3820. ; 51:6, s. 1838-1847
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Journal article (peer-reviewed)abstract
- Steep clinal transitions in one or several inherited characters between genetically distinct populations are usually referred to as hybrid zones. Essentially two different mechanisms may maintain strap genetic dines. Either selection acts against hybrids that are unfit over the entire zone due to their mixed genetic origin (endogenous selection), or hybrids and parental types attain different fitness values in different parts of the dine (exogenous selection). Survival rate estimates of hybrids and parental forms in different regions of the dine may be used to distinguish between these models to assess how the dine is maintained. We used reciprocal transplants to test the relative survival rates of two parental ecotypes and their hybrids over microscale hybrid zones in the direct-developing marine snail Littorina saxatilis (Olivi) on the rocky shores of Galicia, Spain. One of the parental forms occupies upper and the other lower shores, and the hybrids are found at various proportions (1-38%) along with both parental forms in a midshore zone a few meters wide. The survival rate over one month was 39-52% of the native ecotype on upper shores, but only 2-8% for the lower-shore ecotype. In contrast, only 4-8% of the upper-shore ecotype but 53% of large (> 6 mm) and 8% of small (3-6 mm) native lower-shore ecotype survived in the lower shores. In the midshores, both the two parental ecotypes and the hybrids survived about equally well. Thus there is a considerable advantage for the native ecotypes in the upper and lower shores, while in the hybrid zone none of the morphs. hybrids included, are favored. This indicates that the dimorphism of L. saxatilis is maintained by steep cross-shore selection gradients, thus supporting the selection-gradient model of hybrid zones. We performed field and laboratory experiments that suggest physical factors and predation as important selective agents. Earlier studies indicate assortative mating between the two ecotypes in the midshore. This is unexpected in a hybrid zone maintained by selection gradients, and it seems as if the reproductive barrier compresses the hybrid zone considerably.
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