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      Three new species of Carychium O.F. Müller, 1773 from the Southeastern USA, Belize and Panama are described using computer tomography (CT) (Eupulmonata, Ellobioidea, Carychiidae)

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          Abstract

          Abstract

          Three new species of the genus Carychium O.F. Müller, 1773, Carychium hardiei Jochum & Weigand, sp. n., Carychium belizeense Jochum & Weigand, sp. n. and Carychium zarzaae Jochum & Weigand, sp. n. are described from the Southeastern United States, Belize and Panama, respectively. In two consecutive molecular phylogenetic studies of worldwide members of Carychiidae , the North and Central American morphospecies Carychium mexicanum Pilsbry, 1891 and Carychium costaricanum E. von Martens, 1898 were found to consist of several evolutionary lineages. Although the related lineages were found to be molecularly distinct from the two nominal species, the consequential morphological and taxonomic assessment of these lineages is still lacking. In the present paper, the shells of these uncovered Carychium lineages are assessed by comparing them with those of related species, using computer tomography for the first time for this genus. The interior diagnostic characters are emphasized, such as columellar configuration in conjunction with the columellar lamella and their relationship in context of the entire shell. These taxa are morphologically described and formally assigned their own names.

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          Evolution of microgastropods (Ellobioidea, Carychiidae): integrating taxonomic, phylogenetic and evolutionary hypotheses

          Background Current biodiversity patterns are considered largely the result of past climatic and tectonic changes. In an integrative approach, we combine taxonomic and phylogenetic hypotheses to analyze temporal and geographic diversification of epigean (Carychium) and subterranean (Zospeum) evolutionary lineages in Carychiidae (Eupulmonata, Ellobioidea). We explicitly test three hypotheses: 1) morphospecies encompass unrecognized evolutionary lineages, 2) limited dispersal results in a close genetic relationship of geographical proximally distributed taxa and 3) major climatic and tectonic events had an impact on lineage diversification within Carychiidae. Results Initial morphospecies assignments were investigated by different molecular delimitation approaches (threshold, ABGD, GMYC and SP). Despite a conservative delimitation strategy, carychiid morphospecies comprise a great number of unrecognized evolutionary lineages. We attribute this phenomenon to historic underestimation of morphological stasis and phenotypic variability amongst lineages. The first molecular phylogenetic hypothesis for the Carychiidae (based on COI, 16S and H3) reveals Carychium and Zospeum to be reciprocally monophyletic. Geographical proximally distributed lineages are often closely related. The temporal diversification of Carychiidae is best described by a constant rate model of diversification. The evolution of Carychiidae is characterized by relatively few (long distance) colonization events. We find support for an Asian origin of Carychium. Zospeum may have arrived in Europe before extant members of Carychium. Distantly related Carychium clades inhabit a wide spectrum of the available bioclimatic niche and demonstrate considerable niche overlap. Conclusions Carychiid taxonomy is in dire need of revision. An inferred wide distribution and variable phenotype suggest underestimated diversity in Zospeum. Several Carychium morphospecies are results of past taxonomic lumping. By collecting populations at their type locality, molecular investigations are able to link historic morphospecies assignments to their respective evolutionary lineage. We propose that rare founder populations initially colonized a continent or cave system. Subsequent passive dispersal into adjacent areas led to in situ pan-continental or mountain range diversifications. Major environmental changes did not influence carychiid diversification. However, certain molecular delimitation methods indicated a recent decrease in diversification rate. We attribute this decrease to protracted speciation.
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            A new approach to an old conundrum--DNA barcoding sheds new light on phenotypic plasticity and morphological stasis in microsnails (Gastropoda, Pulmonata, Carychiidae).

            The identification of microsnail taxa based on morphological characters is often a time-consuming and inconclusive process. Aspects such as morphological stasis and phenotypic plasticity further complicate their taxonomic designation. In this study, we demonstrate that the application of DNA barcoding can alleviate these problems within the Carychiidae (Gastropoda, Pulmonata). These microsnails are a taxon of the pulmonate lineage and most likely migrated onto land independently of the Stylommatophora clade. Their taxonomical classification is currently based on conchological and anatomical characters only. Despite much confusion about historic species assignments, the Carychiidae can be unambiguously subdivided into two taxa: (i) Zospeum species, which are restricted to karst caves, and (ii) Carychium species, which occur in a broad range of environmental conditions. The implementation of discrete molecular data (COI marker) enabled us to correctly designate 90% of the carychiid microsnails. The remaining cases were probably cryptic Zospeum and Carychium taxa and incipient species, which require further investigation into their species status. Because conventional reliance upon mostly continuous (i.e. nondiscrete) conchological characters is subject to fallibility for many gastropod species assignments, we highly recommend the use of DNA barcoding as a taxonomic, cutting-edge method for delimiting microsnail taxa. © 2010 Blackwell Publishing Ltd.
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              An annotated checklist and bibliography of the land and freshwater snails of México and Central America

              The checklist of the non-marine gastropod fauna summarizes the known species and subspecies that are recognized from México and Central America. It is an annotated list of 1800 terminal taxa presented in a hierarchical framework. These include 1502 native species plus 278 native subspecies, and 20 introduced species. The native species include 175 aquatic operculates, 84 aquatic pulmonates, 130 terrestrial operculates, and 1112 terrestrial pulmonates. In most cases in order to be as objective as possible the list uncritically records the most recent assignment of terminal taxa. In a few instances some changes are deemed necessary. For higher-level changes competing schemes are treated equally. In cases of terminals and higher taxa, readers are directed to the systematic works that discuss relevant taxonomy. It is anticipated that the annotated list will be a useful resource for everyone interested in non-marine gastropods and their nomenclature. In addition to clarifying some issues or points of confusion, this list should also provide an impetus for future work aimed at clarifying and resolving areas of taxonomic disagreement and/or uncertainty, and to make better known the non-marine molluscan fauna of México and Central America.
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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2017
                22 May 2017
                : 675
                : 97-127
                Affiliations
                [1 ] Naturhistorisches Museum der Burgergemeinde Bern, CH-3005 Bern, Switzerland
                [2 ] Institute of Ecology and Evolution, University of Bern, 3012 Bern, Switzerland
                [3 ] Department of Biology, Aquatic Ecosystem Research, University of Duisburg-Essen, 45141 Essen, Germany
                [4 ] Naturhistorisches Museum der Burgergemeinde Bern, CH-3005 Bern, Switzerland
                [5 ] Institute for Ecology, Evolution and Diversity; Senckenberg Biodiversity and Climate Research Center (BiK-F); Senckenberg Gesellschaft für Naturforschung (SGN), Goethe-University (GU), 60438 Frankfurt/M., Germany
                [6 ] Zoologische Staatssammlung München, 81247 München, Germany
                [7 ] Department of Molecular Evolution and Plant Systematics & Herbarium (LZ), Institute of Biology, Leipzig University, 04103 Leipzig, Germany
                [8 ] Department of Experimental Radiology, Justus-Liebig University Giessen, Biomedical Research Center Seltersberg (BFS), 35392 Giessen, Germany
                [9 ] Department of Radiology, Universitätsklinikum Giessen und Marburg GmbH-Standort Giessen, Center for Radiology, 35385 Giessen, Germany
                Author notes
                Corresponding author: Adrienne Jochum (Adrienne.jochum@gmail.com)

                Academic editor: P. Stoev

                Article
                10.3897/zookeys.675.12453
                5523888
                ce2cc121-7047-454b-a0eb-bb94d9abe594
                Adrienne Jochum, Alexander M. Weigand, Estee Bochud, Thomas Inäbnit, Dorian D. Dörge, Bernhard Ruthensteiner, Adrien Favre, Gunhild Martels, Marian Kampschulte

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 27 February 2017
                : 12 April 2017
                Categories
                Research Article

                Animal science & Zoology
                computer tomography,conservation,ecology,leaf litter-dwelling species,microgastropods,shell variability,animalia,eupulmonata,carychiidae

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