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Revision of the Family Alycidae (Acariformes, Acari), with Special Reference to European Species

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Revision of the Family Alycidae (Acariformes, Acari), with Special Reference to European Species Revision of the family Alycidae (Acariformes, Acari), with special reference to European species Matti Uusitalo Department of Biological and Environmental Sciences University of Helsinki Academic Dissertation To be presented for public examination, with permission of the Faculty of Biosciences, in auditorium 6 at B-building (Latokartanonkaari 7) on the 12th of March at 12 noon. Helsinki 2010 Supervisor Dr. Pekka T. Lehtinen University of Turku Finland Rewievers Prof. David Evans Walter University of Alberta Canada Dr. Mark Judson Mus. natl. Hist. nat., Paris France Opponent Dr. Wojciech . Magowski Adam Mickiewicz University Poland ISBN 978-952-92-6869-6 (paperback) ISBN 978-952-10-6096-0 (PDF) http://ethesis.helsinki.fi Helsinki University Print Helsinki 2010 Revision of the family Alycidae (Acariformes, Acari), with special reference to European species Matti Uusitalo Metsälauvalantie 8 I 73, FIN-36220 KANGASALA, Finland Abstract This thesis provides a proposal to divide Alycidae G. Canestrini & Fanzago into two subfamilies and four tribes. Three of the tribes (Alycini G. Canestrini & Fanzago new rank; Bimichaeliini Womersley new rank; Petralycini new rank) include European members and are defined here. Also a preliminary generic division for both the Alycini and Bimichaeliini is introduced. This new hierarchy is based on a reassessment and reranking of new and previously known synapomorphies of the clusters concerned by cladistic analysis, using 60 morphological characters for 48 ingroup, one sister group and one outgroup species. Of special importance are the following conclusions: (1) Petralycini differ from two other major lineages by the unique structure of the prodorsum and palpi; (2) Bimichaeliinae share the presence of large lamellae and the same structure of chelicera, but there are distinct groups on the grounds of secondary and primary patterning of the integument which represent different evolutionary trends; (3) Alycini are united by presence of a reduced pair of lateral eyes but there are distinct groups on the grounds of cheliceral structure and the patterning of integument. The fine morphology of integument was revealed by Scanning Electron Microscopy. The basic characters of the taxa are illustrated either by SEM micrographs or by outline drawings. The subfamilies, tribes, genera and all European species known to now are (re)described and keys to all the ranks are given. The presented classification of Alycini includes the redescriptions of Alycus C.L. Koch, Pachygnathus Dugès and Amphialycus Zachvatkin. Orthacarus Zachvatkin is reduced to a new subgenus of Amphialycus (new rank). The species described or redescribed are: Pachygnathus wasastjernae sp. nov. from Finland; Pachygnathus villosus Dugès [in Oken] (as a senior synonym of P. ornithorhynchus Grandjean); Alycus roseus C.L. Koch; Alycus denasutus (Grandjean) comb. and stat. nov.; Alycus trichotus (Grandjean) comb. nov.; Alycus marinus (Schuster) comb. nov.; Amphialycus (Amphialycus) pentophthalmus Zachvatkin; Amphialycus (Amphialycus) leucogaster (Grandjean); and Amphialycus (Orthacarus) oblongus (Halbert) comb. nov. Alycus rostratus Trägårdh is synonymized with A. roseus; Pachygnathus berlesei Bottazzi and P. lombardinii Bottazzi are synonymized with A. trichotus; Pachygnathus arhinosus Willmann is synonymized with A. leucogaster; and Orthacarus tremli Zachvatkin is synonymized with A. oblongus. The European members of the worldwide Bimichaelia Thor are divided into two genera: Bimichaelia s.st. Thor and Laminamichaelia new genus. Bimichaelia crassipalpis Halbert, B. campylognatha Grandjean and B. rectangula Willmann are synonymized with Bimichaelia sarekensis Trägårdh; Michaelia clavigera nomen nudum in Castagnoli & Pegazzano, Bimichaelia diadema Grandjean and B. praeincisa Willmann are synonymized with Laminamichaelia setigera (Berlese) comb. nov.; Bimichaelia stellaris Womersley and B. ramosa Miheli are synonymized with Laminamichaelia arbusculosa 3 (Grandjean) comb. nov. Bimichaelia augustana (Berlese) and Laminamichaelia subnuda (Berlese) comb. nov. are redescribed. Descriptions of Petralycini and Petralycus unicornis Grandjean are rewritten in accordance with the other diagnoses given here. The Devonian genus Protacarus Hirst was found to share none of the derived character states used in the modern delineation of Alycidae. Coccalicus clavatus Willmann belongs to a tydeoid cluster. Rhyncholophus devius C.L. Koch and Pachygnathus ? cavernicola Oudemans are considered as species inquirendae. The following points are discussed: (1) reasons for the model of description, division into subfamilies and tribes and prodorsal evolution are given; (2) the importance of the sensory organs in defining the species and non-linearity in evolution of the reductive trends of chelicera and palpi between the proposed subfamilies and tribes emphasized; (3) positions of these primitive mites and other major lineages in existing cladograms; and (4) some pros and cons in working with SEM. A database of prodorsa, illustrated in details using SEM, might be an answer to future needs of species identification in soil zoology, ecology and conservation, as proposed in Appendix 2. Selected figures of an undescribed bimichaeliine species ‘sil’ from South Africa, which forms a monotypic sister-group to all the rest of the Bimichaeliinae in the analysis, are given in Appendix 3. Key words: Acari, Endeostigmata, Alycidae, cladistics, classification, new taxa, Bimichaeliinae, Bimichaeliini new rank, Alycinae, Alycini new rank, Petralycini new rank, Europe 4 Table of contents ABSTRACT ................................................................................................................................................ 3 1. INTRODUCTION .................................................................................................................................... 7 1.1. ALYCIDAE G. CANESTRINI & FANZAGO ............................................................................................................. 8 1.1.1. Synonymy ........................................................................................................................................ 8 1.1.2. Characterization of Alycidae ............................................................................................................ 9 1.1.3. Historical background of Alycidae ................................................................................................. 10 1.1.4. Searching for monophyly of Alycidae in the past .......................................................................... 11 2. MATERIAL AND METHODS .................................................................................................................. 13 2.1. THE SPECIMENS ......................................................................................................................................... 13 2.2. PHYLOGENETIC ANALYSIS ............................................................................................................................. 15 2.3. SPECIES INCLUDED ..................................................................................................................................... 16 2.4. CHARACTERS USED ..................................................................................................................................... 17 3. RESULTS ............................................................................................................................................. 27 3.1. RESULTS OF THE PHYLOGENETIC ANALYSIS ....................................................................................................... 27 3.2. KEY TO THE SUBFAMILIES AND TRIBES OF ALYCIDAE ........................................................................................... 32 4. TAXONOMY OF EUROPEAN ALYCINI ................................................................................................... 33 4.1. INTRODUCTION ......................................................................................................................................... 33 4.2. ALYCINI G. CANESTRINI & FANZAGO, 1877 NEW RANK .................................................................................... 34 4.3. KEY TO THE GENERA AND SPECIES OF ALYCINI .................................................................................................. 35 4.4. ALYCUS C.L. KOCH, 1842 ........................................................................................................................... 38 4.4.1. Alycus roseus C.L. Koch, 1842 ........................................................................................................ 40 4.4.2. Alycus denasutus (Grandjean, 1937) comb. and stat. nov. ........................................................... 45 4.4.3. Alycus trichotus (Grandjean, 1937) comb. nov. ............................................................................ 46 4.4.4. Alycus marinus (Schuster, 1958) comb. nov. ................................................................................ 49 4.5. PACHYGNATHUS DUGÈS, 1834 (S.ST.) ........................................................................................................... 50 4.5.1. Pachygnathus villosus Dugès [in Oken], 1836 ............................................................................... 52 4.5.2. Pachygnathus wasastjernae sp. nov. ............................................................................................ 53 4.6. AMPHIALYCUS ZACHVATKIN,
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