02062nas a2200169 4500008004100000022001400041245020500055210006900260260002900329300001400358490000800372520139900380100001701779700001901796700002001815856005701835 1989 eng d a1469-799800aMorphological and biochemical variation in populations of Dugesia (G.) tigrina (Turbellaria, Tricladida, Paludicola) from the western Mediterranean: biogeographical and taxonomical implications0 aMorphological and biochemical variation in populations of iDuges bBlackwell Publishing Ltd a609–6260 v2183 a

Dugesia (Girardia) tigrina is a Nearctic freshwater triclad native to North America, now amply distributed in most European countries as the result of a reportedly unique introduction into north-west Europe early this century and its dispersal southwards and eastwards in the last decades. All European populations described so far are externally spotted, with a pigmented pharynx, and, apart from two sexual populations recently described in England, they are fissiparous. This uniformity contrasts with the variability reported in its native area, thus supporting the hypothesis of a single introduction. However, thorough morphological, karyological and biochemical comparative studies of European populations have never been properly done.We have undertaken such a study comparing 24 populations present in a restricted area of the western Mediterranean. The results show an unexpectedly ample variation in external and pharyngeal pigmentation patterns, the presence of sexually reproducing populations, and ample variation in their biochemical patterns as revealed by isoelectric-focusing. This is an indication that European Dugesia (G.) tigrina is far more diverse than previously thought, suggesting that several, independent introductions from its native area have recently occurred, and the implications of these findings in biogeographical and taxonomic terms are discussed.

1 aRibas, Maria1 aRiutort, Marta1 aBaguñà, Jaume uhttp://dx.doi.org/10.1111/j.1469-7998.1989.tb05003.x00656nas a2200181 4500008004100000245012800041210006900169260001200238300001200250490001400262100002300276700002100299700001700320700001900337700002000356700001900376856007900395 1998 eng d00aA robust molecular phylogeny of the Tricladida (Platyhelminthes: Seriata) with a discussion on morphological synapomorphies0 arobust molecular phylogeny of the Tricladida Platyhelminthes Ser c04/1998 a631-6400 v265(1396)1 aCarranza, Salvador1 aLittlewood, D, T1 aClough, K, A1 aRuiz-Trillo, I1 aBaguñà, Jaume1 aRiutort, Marta uhttp://www.pubmedcentral.nih.gov/picrender.fcgi?artid=1689013&blobtype=pdf00502nas a2200157 4500008004100000245005500041210005500096260002900151300001000180100002000190700002300210700001600233700002400249700001900273856005200292 2001 eng d00aMolecular taxonomy and phylogeny of the Tricladida0 aMolecular taxonomy and phylogeny of the Tricladida aLondonbTaylor & Francis a49-561 aBaguñà, Jaume1 aCarranza, Salvador1 aPaps, Jordi1 aRuiz-Trillo, Iñaki1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/130001957nas a2200157 4500008004100000245012200041210006900163260001200232300001200244490000700256520139900263100002701662700002001689700001901709856007101728 2008 eng d00aMolecular phylogeny of land and freshwater planarians (Tricladida, Platyhelminthes): from freshwater to land and back0 aMolecular phylogeny of land and freshwater planarians Tricladida c05/2008 a555-5680 v473 a

The suborder Tricladida (phylum Platyhelminthes) comprises the well-known free-living flatworms, taxonomically grouped into three infraorders according to their ecology: Maricola (marine planarians), Paludicola (freshwater planarians), and Terricola (land planarians). Molecular analyses have demonstrated that the Paludicola are paraphyletic, the Terricola being the sister group of one of the three paludicolan families, the Dugesiidae. However, neither 18S rDNA nor COI based trees have been able to resolve the relationships among species of Terricola and Dugesiidae, particularly the monophyly of Terricola. Here, we present new molecular data including sequences of nuclear genes (18S rDNA, 28S rDNA) and a mitochondrial gene (COI) of a wider sample of dugesiid and terricolan species. The new sequences have been analyzed, together with those previously obtained, in independent and concatenated analyses using maximum likelihood and bayesian methods. The results show that, although some parts of the trees remain poorly resolved, they support a monophyletic origin for Terricola followed by a likely return of some species to freshwater habitats. Relationships within the monophyletic group of Dugesiidae are clearly resolved, and relationships among some terricolan subfamilies are also clearly established and point to the need for a thorough revision of Terricola taxonomy.

1 aÁlvarez-Presas, Marta1 aBaguñà, Jaume1 aRiutort, Marta uhttp://www.sciencedirect.com/science/article/pii/S105579030800039000577nas a2200157 4500008004100000022001400041245013700055210006900192300001200261490000700273100002000280700002200300700002300322700001900345856005500364 2009 eng d a1463-640900aMolecular analysis of the diversity of terrestrial planarians (Platyhelminthes, Tricladida, Continenticola) in the Iberian Peninsula0 aMolecular analysis of the diversity of terrestrial planarians Pl a637-6490 v381 aMateos, Eduardo1 aCabrera, Cristina1 aCarranza, Salvador1 aRiutort, Marta uhttp://dx.doi.org/10.1111/j.1463-6409.2009.00398.x00582nas a2200145 4500008004100000245008500041210006900126300001600195490000700211100001800218700002400236700001900260700002000279856013700299 2009 eng d00aA new higher classification of planarian flatworms (Platyhelminthes, Tricladida)0 anew higher classification of planarian flatworms Platyhelminthes a1763–17770 v431 aSluys, Ronald1 aKawakatsu, Masaharu1 aRiutort, Marta1 aBaguñà, Jaume uhttp://each.uspnet.usp.br/planarias/Artigos/Geoplanidae/2009-Sluys-et-al-A%20new%20higher%20classification%20of%20planarian%20fl.pdf00526nas a2200145 4500008004100000245011400041210007100155300001000226490000700236100002200243700002700265700001700292700001900309856005200328 2010 eng d00aPlanárias terrestres : Modelo de estudo para a prote{\c c}ão da mata atlântica no corredor da Serra do Mar0 aPlanárias terrestres Modelo de estudo para a prote c c ão da mat a44-490 v451 aCarbayo, Fernando1 aÁlvarez-Presas, Marta1 aRozas, Julio1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/123401912nas a2200193 4500008004100000245018700041210006900228260001200297300001200309490000700321520121200328100002101540700001801561700001601579700003201595700002001627700001901647856005201666 2009 eng d00aMolecular barcoding and phylogeography of sexual and asexual freshwater planarians of the genus Dugesia in the Western Mediterranean (Platyhelminthes, Tricladida, Dugesiidae).0 aMolecular barcoding and phylogeography of sexual and asexual fre c09/2009 a835-8450 v523 a

Planarians of the genus Dugesia have a worldwide distribution with high species diversity in the Mediterranean area. In this area, populations with a triploid karyotype that reproduce by fissiparity are exceptionally frequent, outnumbering the sexual populations. This situation poses interesting questions, such as the age of these asexual lineages, whether they all belong to the same species or whether the triploidization event is recurrent, and what factors (climatic, geographical, historical...) explain the prevalence of these asexual forms. However, asexual populations cannot be assigned to a species due to the lack of copulatory apparatus - the main structure used in species identification. In this study, we have developed a DNA barcoding method, based on COI and ITS-1 sequences, which allows the assignment of the fissiparous forms to sexual species. At the same time, phylogenetic analysis from species of the western Mediterranean have unveiled the presence of species with highly differentiated populations alongside species with a wide distribution and almost no genetic variation. The roles of habitat instability, dispersal capacity and human activities are briefly discussed.

1 aLázaro, Eva, M.1 aSluys, Ronald1 aPala, Maria1 aStocchino, Giacinta, Angela1 aBaguñà, Jaume1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/124100574nas a2200157 4500008004100000245010700041210006900148490000700217100002100224700002600245700002400271700001600295700002000311700001900331856006600350 2011 eng d00aSchmidtea mediterranea phylogeography: an old species surviving on a few Mediterranean islands?0 aiSchmidtea mediterraneai phylogeography an old species surviving0 v111 aLázaro, Eva, M.1 aHarrath, Abdul, Halim1 aStocchino, Giacinta1 aPala, Maria1 aBaguñà, Jaume1 aRiutort, Marta uhttp://www.biomedcentral.com/content/pdf/1471-2148-11-274.pdf00485nas a2200145 4500008004100000245007400041210006900115300001000184490000800194100002700202700002200229700001700251700001900268856005200287 2010 eng d00aMata Atlantica y biodiversidad. Un estudio sobre planarias terrestres0 aMata Atlantica y biodiversidad Un estudio sobre planarias terres a80-820 v2911 aÁlvarez-Presas, Marta1 aCarbayo, Fernando1 aRozas, Julio1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/123001942nas a2200157 4500008004100000245017800041210006900219300001200288490000700300520131100307100002701618700002201645700001701667700001901684856008101703 2011 eng d00aLand planarians (Platyhelminthes) as a model organism for fine-scale phylogeographic studies: understanding patterns of biodiversity in the Brazilian Atlantic Forest hotspot0 aLand planarians Platyhelminthes as a model organism for finescal a887-8960 v243 a

The Brazilian Atlantic Forest is one of the richest biodiversity hotspots of the world. Paleoclimatic models have predicted two large stability regions in its northern and central parts, whereas southern regions might have suffered strong instability during Pleistocene glaciations. Molecular phylogeographic and endemism studies show, nevertheless, contradictory results: although some results validate these predictions, other data suggest that paleoclimatic models fail to predict stable rainforest areas in the south. Most studies, however, have surveyed species with relatively high dispersal rates whereas taxa with lower dispersion capabilities should be better predictors of habitat stability. Here, we have used two land planarian species as model organisms to analyse the patterns and levels of nucleotide diversity on a locality within the Southern Atlantic Forest. We find that both species harbour high levels of genetic variability without exhibiting the molecular footprint of recent colonization or population expansions, suggesting a long-term stability scenario. The results reflect, therefore, that paleoclimatic models may fail to detect refugia in the Southern Atlantic Forest, and that model organisms with low dispersal capability can improve the resolution of these models.

1 aÁlvarez-Presas, Marta1 aCarbayo, Fernando1 aRozas, Julio1 aRiutort, Marta uhttp://onlinelibrary.wiley.com/doi/10.1111/j.1420-9101.2010.02220.x/abstract02235nas a2200169 4500008004100000245011800041210006900159300000900228490000700237520166600244100001901910700002701929700002101956700001801977700001601995856005402011 2012 eng d00aEvolutionary history of the Tricladida and the Platyhelminthes: an up-to-date phylogenetic and systematic account0 aEvolutionary history of the Tricladida and the Platyhelminthes a a5-170 v563 a

Within the free-living platyhelminths, the triclads, or planarians, are the best-known group, largely as a result of long-standing and intensive research on regeneration, pattern formation and Hox gene expression. However, the group’s evolutionary history has been long debated, with controversies ranging from their phyletic structure and position within the Metazoa to the relationships among species within the Tricladida. Over the the last decade, with the advent of molecular phylogenies, some of these issues have begun to be resolved. Here, we present an up-to-date summary of the main phylogenetic changes and novelties with some comments on their evolutionary implications. The phylum has been split into two groups, and the position of the main group (the Rhabdithophora and the Catenulida), close to the Annelida and the Mollusca within the Lophotrochozoa, is now clear. Their internal relationships, although not totally resolved, have been clarified. Tricladida systematics has also experienced a revolution since the implementation of molecular data. The terrestrial planarians have been demonstrated to have emerged from one of the freshwater families, giving a different view of their evolution and greatly altering their classification. The use of molecular data is also facilitating the identification of Tricladida species by DNA barcoding, allowing better knowledge of their distribution and genetic diversity. Finally, molecular phylogenetic and phylogeographical analyses, taking advantage of recent data, are beginning to give a clear picture of the recent history of the Dugesia and Schmidtea species in the Mediterranean.

1 aRiutort, Marta1 aÁlvarez-Presas, Marta1 aLázaro, Eva, M.1 aSolà, Eduard1 aPaps, Jordi uhttp://www.ijdb.ehu.es/web/paper.php?doi=113441mr00627nas a2200169 4500008004100000245016100041210006900202300000900271490000900280100002000289700001800309700001600327700002400343700001900367700001900386856005200405 2012 eng d00aFreshwater planarians (Platyhelminthes, Tricladida) from the Palearctic section of the African continent: new records, with the description of a new species0 aFreshwater planarians Platyhelminthes Tricladida from the Palear a1-150 v31821 aHarrath, A., H.1 aSluys, Ronald1 aMerzoug, D.1 aYacoubi-Khebiza, M.1 aAlwasel, Saleh1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/120602158nas a2200193 4500008004100000245012000041210006900161260001200230300001400242490000800256520150700264100001801771700001801789700002801807700002401835700002001859700001901879856006601898 2013 eng d00aIntegrative delineation of species of Mediterranean freshwater planarians (Platyhelminthes: Tricladida: Dugesiidae)0 aIntegrative delineation of species of Mediterranean freshwater p c11/2013 a523–5470 v1693 a

The paper presents an integrative taxonomic study on dugesiid freshwater flatworms from the north-eastern Mediterranean region by applying both morphological and molecular criteria in the formulation of stable species hypotheses. The morphological information obtained for the specimens was used in a traditional way by comparing the organismal traits of the various populations and candidate species with those of known species, as documented in the taxonomic literature and as revealed by examination of histological sections of museum specimens. In the molecular species delimitation the General Mixed Yule-Coalescent method (GMYC) was used. Results of this study (1) supported the presence of 13 Dugesia species in the Hellenic area (including D. sicula Lepori, 1948, a pan-Mediterranean species), (2) culminated in the description of four new Dugesia species, (3) suggested the presence of two Confirmed Candidate Species, (4) pointed to 12 GMYC-delimited units in Greece and two in Slovakia as Unconfirmed Candidate Species and (5) revealed the presence of an entirely new genus, represented by two newly described species and a third Unconfirmed Candidate Species. Our results revealed a high diversity of dugesiid species in this relatively small region. It is concluded that the morphological features used by taxonomists in comparative studies of dugesiid flatworms generally result in reliable identifications and delineations of species taxa, except in the case of cryptic species.

1 aSluys, Ronald1 aSolà, Eduard1 aGritzalis, Konstantinos1 aVila-Farré, Miquel1 aMateos, Eduardo1 aRiutort, Marta uhttp://onlinelibrary.wiley.com/doi/10.1111/zoj.12077/abstract02974nas a2200121 4500008004100000245010400041210006900145490000700214520252500221100002102746700001902767856006602786 2013 eng d00aDugesia sicula (Platyhelminthes, Tricladida): the colonizing success of an asexual Planarian0 aiDugesia siculai Platyhelminthes Tricladida the colonizing succe0 v133 a

Background Dugesia sicula is the only species of its genus not presenting an endemic or restricted distribution within the Mediterranean area. It mostly comprises fissiparous populations (asexual reproduction by body division and regeneration), most likely sexually sterile, and characterized by an extremely low genetic diversity interpreted as the consequence of a recent anthropic expansion. However, its fissiparous reproduction can result in an apparent lack of diversity within the species, since genetic variation within individuals can be as large as between them because most individuals within a population are clones. We have estimated haplotype and nucleotide diversity of cytochrome oxidase I within and among individuals along the species distribution of a broad sample of D. sicula, including asexual and the two only sexual populations known today; and predicted its potential distribution based on climatic variables. Our aim was to determine the centre of colonisation origin, whether the populations are recent, and whether the species is expanding. Results The species presents 3 most frequent haplotypes, differing in a maximum of 11 base pairs. As expected from their fissiparous mode of reproduction, in half of all the analysed localities many individuals have multiple heteroplasmic haplotypes. The distribution of haplotypes is not geographically structured; however, the distribution of haplotypes and heteroplasmic populations shows higher diversity in the central Mediterranean region. The potential distribution predicted by climatic variables based modelling shows a preference for coastal areas and fits well with the observed data. Conclusions The distribution and frequency of the most frequent haplotypes and the presence of heteroplasmic individuals allow us to gain an understanding of the recent history of the species, together with previous knowledge on its phylogenetic relationships and age: The species most probably originated in Africa and dispersed through the central Mediterranean. After one or multiple populations became triploid and fissiparous, the species colonized the Mediterranean basin, likely both by its own means and helped by human activities. Its present distribution practically fulfils its potential distribution as modelled with climatic variables. Its prevalence in coastal regions with higher water temperatures predicts a likely future expansion to northern and more interior areas following the increase in temperatures due to climate change.

1 aLázaro, Eva, M.1 aRiutort, Marta uhttp://www.biomedcentral.com/content/pdf/1471-2148-13-268.pdf00537nas a2200157 4500008004100000245009400041210006900135260001200204490000600216100002700222700002000249700001900269700002000288700001900308856005200327 2014 eng d00aDiversity of introduced terrestrial flatworms in the Iberian Peninsula: a cautionary tale0 aDiversity of introduced terrestrial flatworms in the Iberian Pen c06/20140 v21 aÁlvarez-Presas, Marta1 aMateos, Eduardo1 aTudó, Àngels1 aJones, Hugh, D.1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/117200452nas a2200121 4500008004100000245010400041210006900145300001200214490000600226100002700232700001900259856005200278 2014 eng d00aPlanarian (Platyhelminthes, Tricladida) Diversity and Molecular Markers: A New View of an Old Group0 aPlanarian Platyhelminthes Tricladida Diversity and Molecular Mar a323-3380 v61 aÁlvarez-Presas, Marta1 aRiutort, Marta uhttps://continenticola.myspecies.info/node/117903182nas a2200277 4500008004100000020001400041245017400055210006900229260001300298300001400311490000700325520228600332653001102618653001702629653000802646653002102654653002602675653001002701653002002711653001902731100001802750700001802768700002802786700001902814856007102833 2013 eng d a1055-790300aFluvial basin history in the northeastern Mediterranean region underlies dispersal and speciation patterns in the genus Dugesia (Platyhelminthes, Tricladida, Dugesiidae)0 aFluvial basin history in the northeastern Mediterranean region u c2013/3// a877 - 8880 v663 a

In this study we analyzed the phylogenetic relationships of eastern Mediterranean freshwater planarians of the genus Dugesia, estimated divergence times for the various clades, and correlated their phylogeographic patterns with geological and paleoclimatic events, in order to discover which evolutionary processes have shaped the present-day distribution of these animals. Specimens were collected from freshwater courses and lakes in continental and insular Greece. Genetic divergences and phylogenetic relationships were inferred by using the mitochondrial gene subunit I of cytochrome oxidase (COI) and the nuclear ribosomal internal transcribed spacer-1 (ITS-1) from 74 newly collected individuals from Greece. Divergence time estimates were obtained under a Bayesian framework, using the COI sequences. Two alternative geological dates for the isolation of Crete from the mainland were tested as calibration points. A clear phylogeographic pattern was present for Dugesia lineages in the Eastern Mediterranean. Morphological data, combined with information on genetic divergences, revealed that eight out of the nine known species were represented in the samples, while additional new, and still undescribed species were detected. Divergence time analyses suggested that Dugesia species became isolated in Crete after the first geological isolation of the island, and that their present distribution in the Eastern Mediterranean has been shaped mainly by vicariant events but also by dispersal. During the Messinian salinity crisis these freshwater planarians apparently were not able to cross the sea barrier between Crete and the mainland, while they probably did disperse between islands in the Aegean Sea. Their dependence on freshwater to survive suggests the presence of contiguous freshwater bodies in those regions. Our results also suggest a major extinction of freshwater planarians on the Peloponnese at the end of the Pliocene, while about 2 Mya ago, when the current Mediterranean climate was established, these Peloponnese populations probably began to disperse again. At the end of the Pliocene or during the Pleistocene, mainland populations of Dugesia colonized the western coast, including the Ionian Islands, which were then part of the continent.

10aAegean10aBiogeography10aCOI10aDivergence times10aFreshwater planarians10aITS-110aMolecular clock10aPhylogeography1 aSolà, Eduard1 aSluys, Ronald1 aGritzalis, Konstantinos1 aRiutort, Marta uhttp://www.sciencedirect.com/science/article/pii/S1055790312004423