Repeated convergent evolution of parthenogenesis in Acariformes (Acari).

The existence of old species-rich parthenogenetic taxa is a conundrum in evolutionary biology. Such taxa point to ancient parthenogenetic radiations resulting in morphologically distinct species. Ancient parthenogenetic taxa have been proposed to exist in bdelloid rotifers, darwinulid ostracods, and in several taxa of acariform mites (Acariformes, Acari), especially in oribatid mites (Oribatida, Acari). Here, we investigate the diversification of Acariformes and their ancestral mode of reproduction using 18S rRNA. Because parthenogenetic taxa tend to be more frequent in phylogenetically old taxa of Acariformes, we sequenced a wide range of members of this taxon, including early-derivative taxa of Prostigmata, Astigmata, Endeostigmata, and Oribatida. Ancestral character state reconstruction indicated that (a) Acariformes as well as Oribatida evolved from a sexual ancestor, (b) the primary mode of reproduction during evolution of Acariformes was sexual; however, species-rich parthenogenetic taxa radiated independently at least four times (in Brachychthonioidea (Oribatida), Enarthronota (Oribatida), and twice in Nothrina (Oribatida), (c) parthenogenesis additionally evolved frequently in species-poor taxa, for example, Tectocepheus, Oppiella, Rostrozetes, Limnozetes, and Atropacarus, and (d) sexual reproduction likely re-evolved at least three times from species-rich parthenogenetic clusters, in Crotonia (Nothrina), in Mesoplophora/Apoplophora (Mesoplophoridae, Enarthronota), and in Sphaerochthonius/Prototritia (Protoplophoridae, Enarthronota). We discuss possible reasons that favored the frequent diversification of parthenogenetic taxa including the continuous long-term availability of dead organic matter resources as well as generalist feeding of species as indicated by natural variations in stable isotope ratios.

Convergent evolution of aquatic life by sexual and parthenogenetic oribatid mites.

Convergent evolution is one of the main drivers of traits and phenotypes in animals and plants. Here, we investigated the minimum number of independent colonisations of marine and freshwater habitats in derived oribatid mites (Brachypylina), a mainly terrestrial taxon. Furthermore, we investigated whether the reproductive mode (sexual vs. thelytokous) is associated with the habitat type (marine, freshwater) where the animals live. We hypothesized that continuous resource availability in freshwater systems fosters asexual reproduction. We used 18S rDNA sequences to construct a molecular phylogeny of oribatid mites from terrestrial, marine and freshwater habitats. The results indicate that aquatic life in oribatid mites evolved at least 3×: once in Limnozetoidea (including only freshwater taxa) and at least twice in Ameronothroidea. In Ameronothroidea the taxon Ameronothridae n. gen. (nr. Aquanothrus) colonized fresh water independently from Selenoribatidae and Fortuyniidae (mainly marine Ameronothroidea). Reproductive mode was associated neither with marine nor with freshwater life; rather, in both habitats sexual and parthenogenetic taxa occur. However, the reproductive mode was related to the stability of the habitat. Species that live underwater permanently tend to be parthenogenetic whereas taxa whose life cycle is often interrupted by flooding, such as marine oribatid mites, or by desiccation, e.g., freshwater-living Ameronothridae n. gen. (nr. Aquanothrus) (Ameronothroidea) species, are mainly sexual, indicating that continuous access to resources indeed favours parthenogenetic reproduction. Findings of our study therefore suggest that parthenogenetic reproduction is not selected for by disturbances but by unlimited access to resources.

Convergent evolution of defense mechanisms in oribatid mites (Acari, Oribatida) shows no "ghosts of predation past".

Oribatid mites are diverse and abundant terrestrial soil arthropods that are involved in decomposition of organic matter and nutrient cycling. As indicated by fossils starting from the Devonian, they evolved varied mechanisms and structures for defense from predators. We investigated four of these defensive structures (ptychoid body, hologastry, mineralization and opisthonotal glands) and used ancestral character state reconstruction to determine whether they evolved convergently and how many times this may have happened. Phylogenetic trees based on 18S rDNA were constructed for 42 oribatid mite species and two outgroup taxa using likelihood and Bayesian algorithms. The results suggest that at least three of the four defensive structures evolved convergently several times; for opisthonotal glands convergent evolution remains equivocal. This high level of convergence indicates that predation has been an important factor throughout the evolution of oribatid mites, contributing to morphological diversity and potentially also to species richness, as there are indications that some taxa radiated after the evolution of defense structures. Despite the ancientness of oribatid mites, defense structures seems to have been rarely lost, suggesting that they still are functional and necessary to reduce predation, rather than being 'ghosts of predation past'.