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      Terrestrial and lacustrine gastropods from the Priabonian (upper Eocene) of the Sultanate of Oman

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          Abstract

          Terrestrial and aquatic gastropods from the upper Eocene (Priabonian) Zalumah Formation in the Salalah region of the Sultanate of Oman are described. The assemblages reflect the composition of the continental mollusc fauna of the Palaeogene of Arabia, which, at that time, formed parts of the southeastern Tethys coast. Several similarities with European faunas are observed at the family level, but are rarer at the genus level. These similarities point to an Eocene (Priabonian) rather than to a Rupelian age, although the latter correlation cannot be entirely excluded. At the species level, the Omani assemblages lack any relations to coeval faunas. This suggests the possible presence of a distinct biogeographic province during the Palaeogene or may simply reflect the extremely sparse non-marine fossil record of the Eocene in the Tethys region. The occurrence of the genera Lanistes, Pila, and Gulella along with some pomatiids, probably related to extant genera, suggests that the modern African–Arabian continental faunas can be partly traced back to Eocene times and reflect very old autochthonous developments. In contrast, the diverse Vidaliellidae went extinct, and the morphologically comparable Neogene Achatinidae may have occupied the equivalent niches in extant environments. Carnevalea Harzhauser and Neubauer nov. gen., Arabiella Kadolsky, Harzhauser and Neubauer nov. gen., Pyrgulella Harzhauser, Kadolsky and Neubauer nov. gen., Salalahia Kadolsky, Harzhauser and Neubauer nov. gen., Omanitopsis Harzhauser and Neubauer nov. gen., Arabicolaria Harzhauser and Neubauer nov. gen., Pacaudiella Harzhauser and Neubauer nov. gen., Goniodomulus Harzhauser and Neubauer nov. gen., Eoquickia Harzhauser and Neubauer nov. gen., Omanillya H. Nordsieck nov. gen. and Omanifera H. Nordsieck nov. gen. are introduced as new genera. Pila neuberti Harzhauser and Neubauer nov. sp., Arabiella arabica Kadolsky, Harzhauser and Neubauer nov. sp., Pyrgulella parva Harzhauser, Kadolsky and Neubauer nov. sp., Salalahia thaytinitiensis Kadolsky, Harzhauser and Neubauer nov. sp., Omanitopsis vandammei Harzhauser and Neubauer nov. sp., Procyclotopsis eocenica Harzhauser and Neubauer nov. sp., Palaeocyclotus kuehschelmi Harzhauser and Neubauer nov. sp., Arabicolaria arabica Harzhauser and Neubauer nov. sp., Pacaudiella omanica Harzhauser and Neubauer nov. sp., Pacaudiella flammulata Harzhauser and Neubauer nov. sp., Goniodomulus solaniformis Harzhauser and Neubauer nov. sp., Cerastus hyznyi Harzhauser and Neubauer nov. sp., Omanillya lunellifera H. Nordsieck nov. sp., Omanillya costellata H. Nordsieck nov. sp., and Omanifera euclista H. Nordsieck nov. sp. are described as new species.

          Kurzfassung

          Terrestrische und aquatische Gastropoden aus der obereozänen (Priabonium) Zalumah-Formation in der Salalah-Region des Sultanats Oman werden beschrieben. Die Vergesellschaftungen reflektieren die Zusammensetzung der kontinentalen Molluskenfaunen des Paläogens Arabiens, das zu dieser Zeit die südöstliche Küste der Tethys formte. Ähnlichkeiten mit europäischen Faunen lassen sich auf Familien- und selten auch auf Gattungsniveau nachweisen. Diese Übereinstimmungen sprechen eher für ein eozänes Alter (Priabonium) als für eine Einstufung ins Rupelium, obwohl letzteres nicht völlig ausgeschlossen werden kann. Auf Artniveau fehlen Bezüge zu gleichalten Faunen. Das könnte auf eine diskrete paläogene Bioprovinz hinweisen oder einfach aus dem extrem dürftigen nicht-marinen Fossilbefund des tethyalen Eozäns resultieren. Das Vorkommen von Gattungen wie Lanistes, Pila und Gulella, gemeinsam mit einigen Pomatiiden, die mit rezenten Gattungen verwandt sein dürften, zeigt, dass die Wurzeln der modernen afrikanisch-arabischen kontinentalen Molluskenfauna zumindest teilweise bis ins Eozän zurückreichen. Im Gegensatz dazu starben die artenreichen Vidaliellidae aus, und die morphologisch ähnlichen Achatinidae dürften ihre ökologischen Nischen im Neogen übernommen haben. Carnevalea Harzhauser und Neubauer nov. gen., Arabiella Kadolsky, Harzhauser und Neubauer nov. gen., Pyrgulella Harzhauser, Kadolsky und Neubauer nov. gen., Salalahia Kadolsky, Harzhauser und Neubauer nov. gen., Omanitopsis Harzhauser und Neubauer nov. gen., Arabicolaria Harzhauser und Neubauer nov. gen., Pacaudiella Harzhauser und Neubauer nov. gen., Goniodomulus Harzhauser und Neubauer nov. gen., Eoquickia Harzhauser und Neubauer nov. gen., Omanillya H. Nordsieck nov. gen. und Omanifera H. Nordsieck nov. gen. Werden als neue Gattungen eingeführt. Pila neuberti Harzhauser und Neubauer nov. sp., Arabiella arabica Kadolsky, Harzhauser und Neubauer nov. sp., Pyrgulella parva Harzhauser, Kadolsky und Neubauer nov. sp., Salalahia thaytinitiensis Kadolsky, Harzhauser und Neubauer nov. sp., Omanitopsis vandammei Harzhauser und Neubauer nov. sp., Procyclotopsis eocenica Harzhauser und Neubauer nov. sp., Palaeocyclotus kuehschelmi Harzhauser und Neubauer nov. sp., Arabicolaria arabica Harzhauser und Neubauer nov. sp., Pacaudiella omanica Harzhauser und Neubauer nov. sp., Pacaudiella flammulata Harzhauser und Neubauer nov. sp., Goniodomulus solaniformis Harzhauser und Neubauer nov. sp., Cerastus hyznyi Harzhauser und Neubauer nov. sp., Omanillya lunellifera H. Nordsieck nov. sp., Omanillya costellata H. Nordsieck nov. sp. und Omanifera euclista H. Nordsieck nov. sp. werden als neue Arten beschrieben.

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          On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia

          Background A robust phylogenetic hypothesis of euthyneuran gastropods, as a basis to reconstructing their evolutionary history, is still hindered by several groups of aberrant, more or less worm-like slugs with unclear phylogenetic relationships. As a traditional "order" in the Opisthobranchia, the Acochlidia have a long history of controversial placements, among others influenced by convergent adaptation to the mainly meiofaunal habitats. The present study includes six out of seven acochlidian families in a comprehensive euthyneuran taxon sampling with special focus on minute, aberrant slugs. Since there is no fossil record of tiny, shell-less gastropods, a molecular clock was used to estimate divergence times within Euthyneura. Results Our multi-locus molecular study confirms Acochlidia in a pulmonate relationship, as sister to Eupulmonata. Previous hypotheses of opisthobranch relations, or of a common origin with other meiofaunal Euthyneura, are clearly rejected. The enigmatic amphibious and insectivorous Aitengidae incerta sedis clusters within Acochlidia, as sister to meiofaunal and brackish Pseudunelidae and limnic Acochlidiidae. Euthyneura, Opisthobranchia and Pulmonata as traditionally defined are non-monophyletic. A relaxed molecular clock approach indicates a late Palaeozoic diversification of Euthyneura and a Mesozoic origin of the major euthyneuran diversity, including Acochlidia. Conclusions The present study shows that the inclusion of small, enigmatic groups is necessary to solve deep-level phylogenetic relationships, and underlines that "pulmonate" and "opisthobranch" phylogeny, respectively, cannot be solved independently from each other. Our phylogenetic hypothesis requires reinvestigation of the traditional classification of Euthyneura: morphological synapomorphies of the traditionally defined Pulmonata and Opisthobranchia are evaluated in light of the presented phylogeny, and a redefinition of major groups is proposed. It is demonstrated that the invasion of the meiofaunal habitat has occurred several times independently in various euthyneuran taxa, leading to convergent adaptations previously misinterpreted as synapomorphies. The inclusion of Acochlidia extends the structural and biological diversity in pulmonates, presenting a remarkable flexibility concerning habitat choice.
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            Freshwater Snails Of Africa And Their Medical Importance

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              The non‐marine aquatic Mollusca of Thailand

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                Author and article information

                Contributors
                mathias.harzhauser@nhm-wien.ac.at
                thomas.neubauer@nhm-wien.ac.at
                kadolsky@btsgeo.com
                pickford@mnhn.fr
                hnords@t-online.de
                Journal
                Palaontol Z
                Palaontol Z
                Palaontologische Zeitschrift
                Springer Berlin Heidelberg (Berlin/Heidelberg )
                0031-0220
                1867-6812
                29 October 2015
                29 October 2015
                2016
                : 90
                : 1
                : 63-99
                Affiliations
                [1 ]Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria
                [2 ]66 Heathhurst Road, Sanderstead, CR2 0BA Surrey, UK
                [3 ]Sorbonne Universités-CR2P, MNHN, CNRS, UPMC-Paris VI, 8, rue Buffon, 75005 Paris, France
                [4 ]Senckenberg Forschungsinstitut Frankfurt; Malakologie, Senckenberganlage 25, 60325 Frankfurt Am Main, Germany
                Article
                277
                10.1007/s12542-015-0277-1
                5448427
                126175ae-6ed3-4c17-8335-40741d01e1cf
                © The Author(s) 2015

                Open AccessThis article is distributed under the terms of the Creative Commons Attribution 4.0 International License ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

                Funding
                Funded by: FundRef http://dx.doi.org/10.13039/501100002428, Austrian Science Fund;
                Award ID: P25365–B25
                Award Recipient :
                Categories
                Research Paper
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                © Paläontologische Gesellschaft 2016

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