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Copepoda, Harpacticoida) and Cletodidae T Zoosyst. Evol. 96 (2) 2020, 455–498 | DOI 10.3897/zse.96.51349 Restructuring the Ancorabolidae Sars (Copepoda, Harpacticoida) and Cletodidae T. Scott, with a new phylogenetic hypothesis regarding the relationships of the Laophontoidea T. Scott, Ancorabolidae and Cletodidae Kai Horst George1 1 Senckenberg am Meer, Abt. Deutsches Zentrum für Marine Biodiversitätsforschung DZMB, Südstrand 44, D-26382 Wilhelmshaven, Germany http://zoobank.org/96C924CA-95E0-4DF2-80A8-2DCBE105416A Corresponding author: Kai Horst George ([email protected]) Academic editor: Kay Van Damme ♦ Received 21 February 2020 ♦ Accepted 10 June 2020 ♦ Published 10 July 2020 Abstract Uncovering the systematics of Copepoda Harpacticoida, the second-most abundant component of the meiobenthos after Nematoda, is of major importance for any further research dedicated especially to ecological and biogeographical approaches. Based on the evolution of the podogennontan first swimming leg, a new phylogenetic concept of the Ancorabolidae Sars and Cletodidae T. Scott sensu Por (Copepoda, Harpacticoida) is presented, using morphological characteristics. It confirms the polyphyletic status of the Ancorabolidae and its subfamily Ancorabolinae Sars and the paraphyletic status of the subfamily Laophontodinae Lang. Moreover, it clarifies the phylogenetic relationships of the so far assigned members of the family. An exhaustive phylogenetic analysis was undertaken using 150 morphological characters, resulting in the establishment of a now well-justified monophylum Ancorabolidae. In that context, the Ancorabolus-lineage sensu Conroy-Dalton and Huys is elevated to sub-family rank. Furthermore, the mem- bership of Ancorabolina George in a rearranged monophylum Laophontodinae is confirmed. Conversely, the Ceratonotus-group sensu Conroy-Dalton is transferred from the hitherto Ancorabolinae to the Cletodidae. Within these, the Ceratonotus-group and its hypothesised sister-group Cletodes Brady are combined to form a monophyletic subfamily Cletodinae T. Scott, subfam. nov. Conse- quently, it was necessary to restructure the Ancorabolidae, Ancorabolinae and Laophontodinae and extend the Cletodidae to include the displacement and exclusion of certain taxa. Moreover, comparison of the Ancorabolidae, Cletodidae, Laophontoidea and other Podogennonta shows that the Ancorabolidae and Cletodidae form sister-groups in a monophylum Cletodoidea Bowman and Abele, which similarly has a sister-group-relationship with the Laophontoidea T. Scott. According to the present study, both taxa constitute a derived monophylum within the Podogennonta Lang. Key Words meiofauna, crustacea, Podogennonta, Cletodoidea, systematics, taxonomy Copyright Kai Horst George. 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. 456 George, K.H.: Restructuring the Ancorabolidae Table of contents Introduction ......................................................................................................................................................................2 Methods............................................................................................................................................................................3 Results ..............................................................................................................................................................................3 Discussion ......................................................................................................................................................................14 I. General remarks .......................................................................................................................................................14 II. The development of the P1 in the Podogennonta as the origin of the new phylogenetic concept ..........................15 III. Phylogenetic analysis: A new concept for a podogennontan Laophontoidea – Ancorabolidae ● – Cletodidae ● -clade, based on the re-evaluation of morphological characters ............................................................17 A. The Laophontoidea–Cletodoidea-clade (L–C-clade) ...........................................................................................17 B. Cletodoidea Bowman & Abele, 1982 ...................................................................................................................18 C. Ancorabolidae ● Sars, 1909 ..................................................................................................................................20 D. Laophontodinae ● Lang, 1944 .............................................................................................................................21 E. Ancorabolinae ● Sars, 1909 ..................................................................................................................................24 F. The Algensiella–Probosciphontodes-clade (A–P-clade) .......................................................................................27 G. Tapholaophontodes ...............................................................................................................................................28 H. The Calypsophontodes–Probosciphontodes-clade (C–P-clade) ...........................................................................28 I. Algensiella ..............................................................................................................................................................29 J. The Laophontodes–Probosciphontodes-clade (La–P-clade) .................................................................................29 K. Calypsophontodes .................................................................................................................................................29 L. The Rostrophontodes–Ancorabolina-clade (R–A-clade) ......................................................................................30 M. Rostrophontodes ..................................................................................................................................................31 N. The Laophontodes–Ancorabolina-clade (La–A-clade) ........................................................................................32 O. Laophontodes .......................................................................................................................................................32 P. The Bicorniphontodes–Ancorabolina-clade (B–A-clade) .....................................................................................33 Q. Bicorniphontodes ..................................................................................................................................................33 R. Ancorabolina ........................................................................................................................................................34 S. The Paralaophontodes–Probosciphontodes-clade (Pa–P-clade) ..........................................................................34 T. Paralaophontodes .................................................................................................................................................35 U. The Lobopleura–Probosciphontodes-clade (Lo–P-clade) ....................................................................................35 V. Lobopleura ............................................................................................................................................................35 W. Probosciphontodes ...............................................................................................................................................36 X. Cletodidae ● T. Scott, 1904 ..................................................................................................................................36 Y. Cletodinae T. Scott, 1904, subfam. nov. ...............................................................................................................36 Z. Remaining Cletodidae ● .......................................................................................................................................37 AA. The Ceratonotus-group (C-g) ............................................................................................................................37 BB. Cletodes ..............................................................................................................................................................39 CC. The status of Patagoniaella Pallares, 1968 ........................................................................................................39 Conclusions ....................................................................................................................................................................40 Acknowledgements ........................................................................................................................................................41 References ......................................................................................................................................................................41 Introduction With an estimated number of > 50,000 species (cf. George et al. 2020 and references therein), the Copepo- Um dahin zu gelangen, muß man sich indessen von da (Crustacea)
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