Arachnida, Solifugae) with Special Focus on Functional Analyses and Phylogenetic Interpretations
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HISTOLOGY AND ULTRASTRUCTURE OF SOLIFUGES Comparative studies of organ systems of solifuges (Arachnida, Solifugae) with special focus on functional analyses and phylogenetic interpretations HISTOLOGIE UND ULTRASTRUKTUR DER SOLIFUGEN Vergleichende Studien an Organsystemen der Solifugen (Arachnida, Solifugae) mit Schwerpunkt auf funktionellen Analysen und phylogenetischen Interpretationen I N A U G U R A L D I S S E R T A T I O N zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) an der Mathematisch-Naturwissenschaftlichen Fakultät der Ernst-Moritz-Arndt-Universität Greifswald vorgelegt von Anja Elisabeth Klann geboren am 28.November 1976 in Bremen Greifswald, den 04.06.2009 Dekan ........................................................................................................Prof. Dr. Klaus Fesser Prof. Dr. Dr. h.c. Gerd Alberti Erster Gutachter .......................................................................................... Zweiter Gutachter ........................................................................................Prof. Dr. Romano Dallai Tag der Promotion ........................................................................................15.09.2009 Content Summary ..........................................................................................1 Zusammenfassung ..........................................................................5 Acknowledgments ..........................................................................9 1. Introduction ............................................................................. 12 1.1. BIOLOGY, PHYSIOLOGY AND LIFE CYCLE ...................................... 12 1.2. GROSSMORPHOLOGY ................................................................ 13 1.3. DISTRIBUTION, SYSTEMATICS AND PHYLOGENY ............................ 16 1.4. OBJECTIVES OF THE PRESENT WORK .......................................... 20 2. Materials and methods ............................................................21 2.1. MATERIALS .............................................................................. 21 2.1.1. Animals ............................................................................... 21 2.1.2. Collecting of animals ............................................................... 22 2.1.3. Chemicals and solutions ........................................................... 25 2.2. METHODS ............................................................................... 31 2.2.1. Keeping live animals ............................................................... 31 2.2.2. Identification ........................................................................ 31 2.2.3. Light microscopy .................................................................... 33 2.2.4. Electron microscopy ............................................................... 36 2.2.5. Stereo microscopy and digital image processing ............................... 37 2.2.6. Video recording ..................................................................... 37 2.2.7. DNA extraction, PCR and Sequencing ............................................ 38 3. Results and discussions ........................................................... 39 3.1. FLUORESCENCE ....................................................................... 39 3.1.1. Results ................................................................................ 39 3.1.2. Discussion ............................................................................ 39 3.2. SUCTORIAL ORGAN .................................................................. 41 3.2.1. Results ................................................................................ 41 External morphology of the suctorial organ ............................................... 41 Histology of the suctorial organ ............................................................ 43 Content Fine structure of the pad .................................................................. 43 3.2.2. Discussion ............................................................................ 43 3.3. TARSALIA OF LEGS I-IV .............................................................. 50 3.3.1. Results ................................................................................ 50 3.3.1.1. Ammotrechidae ................................................................... 50 3.3.1.2. Daesiidae .......................................................................... 55 3.3.1.3. Eremobatidae ..................................................................... 57 3.3.1.4. Galeodidae ......................................................................... 63 3.3.1.5. Gylippidae ........................................................................ 67 3.3.1.6. Solpugidae ........................................................................ 68 3.3.2. Discussion ............................................................................ 70 3.4. MEDIAN EYES .......................................................................... 73 3.4.1. Results ................................................................................ 73 Gross morphology of the ocular tubercle .................................................. 73 Histology of the median eyes .............................................................. 75 Fine structure of the median eyes ......................................................... 75 3.4.2. Discussion ............................................................................ 79 3.5. CENTRAL NERVOUS SYSTEM (CNS) .............................................. 82 3.5.1. Results ................................................................................ 82 Prosomal ganglion .......................................................................... 82 Opisthosomal ganglion ..................................................................... 82 Peripheral nerves .......................................................................... 88 3.5.2. Discussion ............................................................................ 88 3.6. MALLEOLI ............................................................................... 90 3.6.1. Results ................................................................................ 90 3.6.2. Discussion ............................................................................ 94 3.7. ALIMENTARY SYSTEM ................................................................ 96 3.7.1. Results ................................................................................ 96 External morphology and histology of the chelicerae ..................................... 96 External morphology of the rostrum ........................................................ 96 Histology of the rostrum ................................................................... 96 Gut system ................................................................................. 99 Parasites ................................................................................... 111 3.7.2. Discussion ...........................................................................112 Content 3.8. COXAL GLANDS ....................................................................... 116 3.8.1. Results ............................................................................... 116 Anatomy of the coxal glands .............................................................. 116 Histology of the coxal glands ............................................................... 116 Fine structure of the coxal glands ........................................................ 120 3.8.2. Discussion ...........................................................................120 3.9. RESPIRATORY SYSTEM .............................................................. 124 3.9.1. Results ............................................................................... 124 Prosomal spiracles and tracheae ..........................................................124 Tracheal cells and tracheae ............................................................... 127 Opisthosomal spiracles ....................................................................127 3.9.2. Discussion ...........................................................................127 Spiracles ................................................................................... 127 Tracheae .................................................................................. 129 3.10. CIRCULATORY SYSTEM ............................................................ 132 3.10.1. Results ............................................................................. 132 Heart ...................................................................................... 132 Hemocytes ................................................................................ 132 Nephrocytes ...............................................................................138 3.10.2. Discussion ......................................................................... 139 3.11. FEMALE GENITAL SYSTEM ....................................................... 142 3.11.1. Results ............................................................................. 142 Juvenile female genital system ........................................................... 143 Adult female genital system .............................................................. 148 Ultrastructure of the adult female genital system ........................................151