Revision of Trogulidae Sundevall, 1833 (Arachnida: Opiliones) Dissertation zur Erlangung des Grades „Doktor der Naturwissenschaften“ Am Fachbereich Biologie Der Johannes Gutenberg-Universität Mainz Axel Ludwig Schönhofer geboren am 3. Januar 1976 in Mainz Mainz, 17. 4. 2009 Mainzer Dissertation - D77 1 Dekan: Erstberichterstatter: Zweitberichterstatter: Tag der Prüfung: 30.7.2009 2 To go where no man’s ever been, to find what no mans eye has ever seen, to bring this secret to the world and share, for: “Yes, I am Biologist!” “Der Schmetterlingsjäger” Carl Spitzweg (1840) To my family and friends. For you I am thoroughly thankful! 3 The chapter 4 has been accepted for publication and the chapter 5 has been published in a slightly modified version: Chapter 4: SCHÖNHOFER, A.L., MARTENS J. (in press): Revision of the genus Trogulus Latreille: the Trogulus hirtus species-group (Opiliones: Trogulidae). In Kropf, C. & Horak, P. (eds). Towards a natural history of arthropods and other organisms. In memoriam Konrad Thaler. Contributions to Natural History 12 (accepted 4.5.2007). Species epithets to be used for the decription of new species in the published version of this chapter are here addressed as “manuscript name” [man.n.] prior to publication. Chapter 5: SCHÖNHOFER, A.L., MARTENS, J. (2008): Revision of the genus Trogulus Latreille: the Trogulus coriziformis species-group of the western Mediterranean (Opiliones: Trogulidae). Invertebrate Systematics 22 (5), 523-554. Four species included in this chapter have been published in 2008. In this chapter I uphold “sp. nov.”. In the other chapters the authors are annotated. 4 Contents 1. Abstract ................................................................................................................................ 7 2. General Introduction .......................................................................................................... 8 3. Material and Method ........................................................................................................ 10 3.1. Material ................................................................................................................ 10 3.2. Molecular Genetic ................................................................................................ 11 3.2.1. Laboratory protocols ............................................................................. 11 3.2.2. Phylogenetic analysis ............................................................................ 13 3.3. Morphological examination and morphometry .................................................... 14 3.4. Processing of distributional data .......................................................................... 17 4. Revision of the genus Trogulus Latreille: the Trogulus hirtus species-group (Opiliones: Trogulidae) .........................................................................................................................18 4.1. Introduction ................................................................................................................. 18 4.2. Material ....................................................................................................................... 19 4.3. Results ......................................................................................................................... 20 4.4. Discussion ................................................................................................................... 24 4.5. Taxonomy .................................................................................................................... 26 4.5.1. The Trogulus hirtus species-group ...................................................................... 26 4.5.2. Key to the species of the Trogulus hirtus species-group ..................................... 29 4.5.3. Species accounts .................................................................................................. 31 4.5.3.1. The Trogulus hirtus clade ................................................................................. 31 4.5.3.2. The Trogulus falcipenis clade ........................................................................... 47 4.6. Abstract ....................................................................................................................... 60 5. Revision of the genus Trogulus Latreille: the Trogulus coriziformis species-group of the western Mediterranean (Opiliones: Trogulidae) .................................................... 61 5.1. Introduction ................................................................................................................. 61 5.2. Material ....................................................................................................................... 62 5.3. Results ......................................................................................................................... 64 5.4. Discussion ................................................................................................................... 70 5.4.1. Delineation of species-groups ............................................................................. 70 5.4.2. Phylogenetic relationships within the group ....................................................... 71 5.4.3. Biogeography ....................................................................................................... 72 5.5. Taxonomy .................................................................................................................... 73 5.5.1. Diagnosis of the Trogulus coriziformis species-group ........................................ 73 5.5.2. Key to species of the Trogulus coriziformis species-group ................................. 75 5.5.3. Species accounts .................................................................................................. 77 5.6. Abstract ..................................................................................................................... 114 5 6. Hidden Mediterranean diversity: assessing species taxa by molecular phylogeny within the opilionid family Trogulidae (Arachnida, Opiliones) ................................. 116 6.1. Introduction ............................................................................................................... 116 6.1.1. Present family-level systematics ....................................................................... 116 6.1.2. Present systematics of Trogulidae ..................................................................... 117 6.1.3. Species delineation and cryptic diversity .......................................................... 117 6.2. Material ..................................................................................................................... 118 6.3. Results ....................................................................................................................... 119 6.4. Discussion ................................................................................................................. 129 6.4.1. Taxonomic inference on family level ................................................................ 129 6.4.2. Trogulidae: Taxonomic inference on genus and subfamily .............................. 129 6.4.3. Systematics of Trogulus .................................................................................... 130 6.4.4. Species delineation in Trogulus ......................................................................... 134 6.4.5. Cryptic diversity within Trogulidae .................................................................. 136 6.4.6. Nemastomatidae: Implications from a reduced taxon sampling ...................... 137 6.5. Abstract ..................................................................................................................... 138 7. Distinctive features of Trogulidae genetics ................................................................... 140 7.1. Introduction .............................................................................................................. 140 7.2. Material .................................................................................................................... 140 7.3. Results ...................................................................................................................... 141 7.3.1. Overlay of sequences ......................................................................................... 141 7.3.2. Ambiguous base positions ................................................................................. 142 7.3.3. Diversity in clones from single individuals ....................................................... 145 7.3.4. Unexpected long fragments of ND1 PCR .......................................................... 146 7.4. Discussion ................................................................................................................ 147 7.4.1. Heteroplasmy ..................................................................................................... 147 7.4.2. Possibility of gene rearrangements .................................................................... 148 7.4.3. Effects of micro-organisms on mtDNA ............................................................. 149 7.4.4. Usability of the cytochrome b gene as molecular marker for Opiliones............ 149 8. General conclusions .......................................................................................................