The freshwater flatworm genus Schmidtea is endemic in the Western Palearctic region, where it is represented by only four species, thus contrasting with the high species diversity of the closely related genus Dugesia within Europe. Although containing an important model species in developmental and regeneration research, viz. Schmidtea mediterranea, no evolutionary studies on the genus Schmidtea have been undertaken. For the first time, we present a well‐resolved molecular phylogenetic tree of the four species of the genus, inferred on the basis of two molecular markers, and provide also the first detailed morphological account of Schmidtea nova. The phylogenetic tree generated corroborates an earlier speciation hypothesis based on karyological data and points to chromosomal rearrangements as the main drivers of speciation in this genus. The high genetic divergence between the four species, in combination with previous dating studies and their current geographic distribution, suggests that Schmidtea could have originated in Laurasia but lost most of its diversity during the Oligocene. Thus, its present distribution pattern may be the result of the expansion of three of its four relictual species over Europe, probably after the Pleistocene glaciations. Our detailed morphological study of S. nova revealed that it shows a number of remarkable features: interconnected testis follicles, parovaria, an ejaculatory duct exiting into the primary as well as the secondary seminal vesicle by means of a nipple, and the wall of the distal section of the ejaculatory duct being sclerotic or chitinized.

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Background Predicted genetic consequences of asexuality include high intraindividual genetic diversity (i.e., the Meselson effect) and accumulation of deleterious mutations (i.e., Muller’s Ratchet), among others. These consequences have been largely studied in parthenogenetic organisms, but studies on fissiparous species are scarce. Differing from parthenogens, fissiparous organisms inherit part of the soma of the progenitor, including somatic mutations. Thus, in the long term, fissiparous reproduction may also result in genetic mosaicism, besides the presence of the Meselson effect and Muller’s Ratchet. Dugesiidae planarians show outstanding regeneration capabilities, allowing them to naturally reproduce by fission, either strictly or combined with sex (facultative). Therefore, they are an ideal model to analyze the genetic footprint of fissiparous reproduction, both when it is alternated with sex and when it is the only mode of reproduction. Results In the present study, we generate and analyze intraindividual cloned data of a nuclear and a mitochondrial gene of sexual, fissiparous and facultative wild populations of the species Dugesia subtentaculata . We find that most individuals, independently of their reproductive strategy, are mosaics. However, the intraindividual haplotype and nucleotide diversity of fissiparous and facultative individuals is significantly higher than in sexual individuals, with no signs of Muller’s Ratchet. Finally, we also find that this high intraindividual genetic diversity of fissiparous and facultative individuals is composed by different combinations of ancestral and derived haplotypes of the species. Conclusions The intraindividual analyses of genetic diversity point out that fissiparous reproduction leaves a very special genetic footprint in individuals, characterized by mosaicism combined with the Meselson effect (named in the present study as the mosaic Meselson effect ). Interestingly, the different intraindividual combinations of ancestral and derivate genetic diversity indicate that haplotypes generated during periods of fissiparous reproduction can be also transmitted to the progeny through sexual events, resulting in offspring showing a wide range of genetic diversity and putatively allowing purifying selection to act at both intraindividual and individual level. Further investigations, using Dugesia planarians as model organisms, would be of great value to delve into this new model of genetic evolution by the combination of fission and sex. Electronic supplementary material The online version of this article (10.1186/s12862-019-1440-1) contains supplementary material, which is available to authorized users.

Aim Species biogeography mainly focuses on palaeogeographical events, while environmental factors are generally overlooked despite their importance in species diversification. Here, we use an integrative approach to understand how palaeogeographical and environmental processes shape species distribution and focus on freshwater planarians as the model system. Location Western Mediterranean. Taxon Dugesia. Methods We inferred the phylogenetic relationships of most known Dugesia species in the area using six molecular markers. We then estimated their divergence times and reconstructed their ancestral distribution ranges. We also performed environmental niche modelling analyses using Dugesia subtentaculata as a model to evaluate the effects of several hydro‐environmental variables and the likely existence of interspecific competition on Dugesia distributions. Results Our results provide a new phylogenetic scheme for Dugesia from the Western Mediterranean and show that the time splits between the lineages and their putative ancestral distribution ranges are correlated with microplate tectonic dynamics within the region during the Oligocene–Miocene period. Our environmental niche modelling analyses indicate that the type of land cover and the slope of the terrain are the most important abiotic factors driving the distribution of Dugesia from this region. Finally, we found a partial niche overlap between D. subtentaculata and two other common planarian species from the Iberian Peninsula. Main conclusions The microplate tectonic dynamics of the Western Mediterranean during the Oligocene–Miocene period, together with the position of the mountain ranges and posterior climate changes, may have played crucial roles in driving the biogeographical history of Dugesia in this region. Moreover, both interspecific competition and changes in fluvial characteristics driven by human activities may affect the current diversity and distribution of Dugesia in the Western Mediterranean. This study highlights the importance of integrating different types of information to study the biogeographical history of a species.

Aim Freshwater planarians may have a wide geographical range despite their assumed low vagility. Found across four continents, Dugesia may have either an ancient origin on a large palaeo landmass, followed by colonisation in different regions before continental fragmentation, or a more recent origin and subsequent transoceanic dispersal. We seek to resolve between these two hypotheses. Location Africa, Eurasia and Australasia. Taxon Genus Dugesia (Platyhelminthes: Tricladida: Dugesiidae). Methods We used data from the sequencing of six gene fragments and comprehensive taxonomic sampling of Dugesia from across its distribution range to reconstruct the phylogeny of this genus using maximum likelihood and bayesian inference methods. We conducted two phylogenetic dating analyses using Platyhelminthes fossils and palaeogeological events. Basing on the time‐calibrated molecular phylogenetic framework we evaluated the contribution of vicariance and dispersal to the biogeographical evolution of Dugesia. By reconstructing the ancestral areas and present‐day potential distribution using BioGeoBEARS and niche modelling, we elucidated the biogeographical history of the genus. Results The present‐day distribution of Dugesia is a result of different vicariance and dispersal events. However, we also found evidence of transoceanic dispersal. Consistent with previous hypotheses, Dugesia dates to the Upper Jurassic in the Afro‐Malagasy Gondwana region. We unveiled a novel biogeographical scenario for the genus, involving multiple events of colonisation in Eurasia from continental Africa via at least three dispersal routes. Main conclusions Dugesia is an ancient genus having reached its present distribution through a complex history of dispersal and vicariant events following its origin in southern Gondwana. Despite the low vagility of Dugesia, we found evidence of their overseas dispersal.