Complex evolutionary processes maintain an ancient chromosomal inversion.

Genome re-arrangements such as chromosomal inversions are often involved in adaptation. As such, they experience natural selection, which can erode genetic variation. Thus, whether and how inversions can remain polymorphic for extended periods of time remains debated. Here we combine genomics, experiments, and evolutionary modeling to elucidate the processes maintaining an inversion polymorphism associated with the use of a challenging host plant (Redwood trees) in Timema stick insects. We show that the inversion is maintained by a combination of processes, finding roles for life-history trade-offs, heterozygote advantage, local adaptation to different hosts, and gene flow. We use models to show how such multi-layered regimes of balancing selection and gene flow provide resilience to help buffer populations against the loss of genetic variation, maintaining the potential for future evolution. We further show that the inversion polymorphism has persisted for millions of years and is not a result of recent introgression. We thus find that rather than being a nuisance, the complex interplay of evolutionary processes provides a mechanism for the long-term maintenance of genetic variation.

Evolution of Trichobaris (Curculionidae) in relation to host plants: Geometric morphometrics, phylogeny and phylogeography.

The family Curculionidae (Coleoptera), the "true" weevils, have diversified tightly linked to the evolution of flowering plants. Here, we aim to assess diversification at a lower taxonomic level. We analyze the evolution of the genus Trichobaris in association with their host plants. Trichobaris comprises eight to thirteen species; their larvae feed inside the fruits of Datura spp. or inside the stem of wild and cultivated species of Solanaceae, such as potato, tobacco and tomato. We ask the following questions: (1) does the rostrum of Trichobaris species evolve according to the plant tissue used to oviposit, i.e., shorter rostrum to dig in stems and longer to dig in fruits? and (2) does Trichobaris diversify mainly in relation to the use of Datura species? For the first question, we estimated the phylogeny of Trichobaris based on four gene sequences (nuclear 18S and 28S rRNA genes and mitochondrial 16S rRNA and COI genes). Then, we carried out morphogeometric analyses of the Trichobaris species using 75 landmarks. For the second question, we calibrated a COI haplotype phylogeny using a constant rate of divergence to infer the diversification time of Trichobaris species, and we traced the host plant species on the haplotype network. We performed an ancestral state reconstruction analysis to infer recent colonization events and conserved associations with host plant species. We found that ancestral species in the Trichobaris phylogeny use the stem of Solanum plants for oviposition and display weak sexual dimorphism of rostrum size, whereas other, more recent species of Trichobaris display sexual dimorphism in rostrum size and use the fruits of Datura species, and a possible reversion to use the stem of Solanaceae was detected in one Trichobaris species. The use of Datura species by Trichobaris species is widely distributed on haplotype networks and restricted to Trichobaris species that originated ca. 5 ±â€¯1.5 Ma. Given that the origin of Trichobaris is estimated to be ca. 6 ±â€¯1.5 Ma, it is likely that Datura has played a role in its diversification.

Phylogeography of specialist weevil Trichobaris soror: a seed predator of Datura stramonium.

Can the genetic structure of a specialist weevil be explained by the geological history of their distribution zone? We analyze the genetic variation of the weevil Trichobaris soror, a specialist seed predator of Datura stramonium, in order to address this question. For the phylogeographic analysis we used the COI gene, and assessed species identity in weevil populations through geometric morphometric approach. In total, we found 53 haplotypes in 413 samples, whose genetic variation supports the formation of three groups: (1) the Transmexican Volcanic Belt (TVB group), (2) the Sierra Madre Sur (SMS group) and (3) the Balsas Basin (BB group). The morphometric analysis suggests that BB group is probably not T. soror. Our results have two implications: first, the phylogeographic pattern of T. soror is explained by both the formation of the geological provinces where it is currently distributed and the coevolution with its host plant, because the TVB and SMS groups could be separated due to the discontinuity of altitude between the geological provinces, but the recent population expansion of TVB group and the high frequency of only one haplotype can be due to specialization to the host plant. Second, we report a new record of a different species of weevil in BB group parasitizing D. stramonium fruits.

Helmet Shape and Phylogeography of the Treehopper Membracis mexicana.

Membracis mexicana (Hemiptera: Membracidae) is distributed in four biogeographic provinces of Mexico. Field observations indicate that there are different forms of this species, but the distribution of the phenotype and the genetic variation of this species have not been clarified. The aim of this study was to quantify the phenotypic and genetic variation of M. mexicana and determine whether the configuration of biogeographic provinces impacts the distribution of this variation. To achieve this, we analyzed 307 photographs using 19 landmarks and geometric morphometrics to quantify the phenotypic variation in helmets. We sequenced five molecular markers for 205 individuals to describe the phylogeographic pattern. As a result, we identified three morphological configurations of the helmet of M. mexicana and two genetic lineages. The morphotypes are (1) a large and wide helmet with small dorsal spots, (2) a small and narrow helmet with large dorsal spots, and (3) a small and narrow helmet with small spots. Genetic lineages are distributed in southeast and western Mexico. The western lineage corresponds to two helmet morphotypes (1 and 2) and the southeast lineage to morphotype 3. We found that the larger helmets correspond to the western lineage and are distributed in Trans-Mexican Volcanic Belt and Pacific lowlands provinces, whereas the smallest helmets correspond to the southeast lineage and are present in the Veracruzan and Yucatan Peninsula provinces.

Large-scale mutation in the evolution of a gene complex for cryptic coloration.

The types of mutations affecting adaptation in the wild are only beginning to be understood. In particular, whether structural changes shape adaptation by suppressing recombination or by creating new mutations is unresolved. Here, we show that multiple linked but recombining loci underlie cryptic color morphs of Timema chumash stick insects. In a related species, these loci are found in a region of suppressed recombination, forming a supergene. However, in seven species of Timema, we found that a megabase-size "supermutation" has deleted color loci in green morphs. Moreover, we found that balancing selection likely contributes more to maintaining this mutation than does introgression. Our results show how suppressed recombination and large-scale mutation can help to package gene complexes into discrete units of diversity such as morphs, ecotypes, or species.