Histology and ultrastructure of solifuges [Elektronische Ressource] : comparative studies of organ systems of solifuges (Arachnida, Solifugae) with special focus on functional analyses and phylogenetic interpretations = Histologie und Ultrastruktur der Solifugen / vorgelegt von Anja Elisabeth Klann

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 04.06.2009Prof. Dr. Klaus FesserDekan ........................................................................................................
Prof. Dr. Dr. h.c. Gerd Alberti
Erster Gutachter ..........................................................................................
Prof. Dr. Romano Dallai
Zweiter Gutachter ........................................................................................
15.09.2009Tag der Promotion 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 ............................................................ 43Content
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 ........................................................................... 112Content
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
Insemination .............................................................................. 155
3.11.2. Discussion ......................................................................... 160
3.12. MALE REPRODUCTIVE SYSTEM ................................................. 166
3.12.1. Results 166
Flagella .................................................................................... 166
General anatomy and histology of the genital system .................................... 166
Ultrastructure of the male genital system ................................................ 170
Spermatozoa 174
3.12.1.1. Ammotrechidae ........................................................