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Abstract
Solenogastres (Mollusca) have a quite uniform bodyplan and an evolutionary history with few shifts out of their deep-water habitat and beyond their epibenthic lifestyle. Consequently, few clades inhabit the shallow subtidal mesopsammon; only Meiomeniidae (order Pholidoskepia) is entirely restricted to this habitat. What was initially designed as a comparative microanatomical redescription of Meiomeniidae to explore the diversity of this clade with its unique evolution, developed into a taxonomic nightmare of cryptic, co-occurring lineages: three out of four valid species of Meiomeniidae co-occur in coarse sands in the Bermuda archipelago and were re-collected at the respective type localities. We analyzed the material combining three-dimensional (3D) reconstructions from histological serial sections and ultrastructural data, providing novel insights into meiomeniid anatomy and discussing potential phylogenetic implications. However, not all collected material could be unambiguously assigned to known lineages of mesopsammic Solenogastres. In addition to meiomeniids, we discovered another co-occurring, externally highly cryptic but anatomically distinguishable lineage. It is provisionally placed within Dondersiidae, but its taxonomic assignment remains problematic due to an exclusive character mosaic and a unique foregut gland complex. Our study reveals the risk of creating chimeric taxa in small-bodied Solenogastres, as morphological characters needed for species delineation cannot be extracted from single individuals, while conspecifity based on external features is risky to assume with cryptic species co-occurring. Molecular markers will be needed to reliably retrieve Meiomeniidae from their current Bermuda Triangle of taxonomy and to proceed in solenogaster taxonomy confronted with a wealth of poorly known lineages especially in meiofaunal forms.
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Acknowledgments
Our special thanks go to Dr. Jon Norenburg (Smithsonian Institution), who introduced KMJ and FSB to meiofaunal Solenogastres and highlighted the need for taxonomic revisions. We also would like to thank Dr. Kevin M. Kocot (University of Alabama) for valuable discussion on meiomeniid evolution and for providingM. swedmarki collected at the type locality. Dr. Alexander Fahrner and Dr. Michael Schrödl (both ZSM) contributed in collecting specimens in Bermuda. The late Alenka Kerin (LMU) assisted with embedding and ultrathin sectioning of the meiomeniid material collected in 1999, Eva Lodde-Bensch (ZSM) sectioned two of the histological semithin series, and Heidemarie Gensler (LMU) helped with additional lab work. Thanks also to Dr. Joachim Haug (LMU) for an introduction to birefringence microscopy. This study was funded by a mini-Advancing Revisionary Taxonomy and Systematics (ARTS) grant of the Society of Systematic Biologists to FSB. We thank two anonymous reviewers for their helpful comments which improved the manuscript.
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Department Biology II, Ludwig-Maximilians-University, Großhaderner Str. 2, 82152, Planegg-Martinsried, Germany
Franziska S. Bergmeier, Gerhard Haszprunar & Katharina M. Jörger
SNSB-Bavarian State Collection of Zoology, Münchhausenstr. 21, 81247, Munich, Germany
Gerhard Haszprunar & Katharina M. Jörger
University Museum of Bergen, Allégaten 41, 5007, Bergen, Norway
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Bergmeier, F.S., Haszprunar, G., Todt, C.et al. Lost in a taxonomic Bermuda Triangle: comparative 3D-microanatomy of cryptic mesopsammic Solenogastres (Mollusca).Org Divers Evol16, 613–639 (2016). https://doi.org/10.1007/s13127-016-0266-6
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