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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

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236 pages
HISTOLOGY AND ULTRASTRUCTURE OF SOLIFUGESComparative studies of organ systems of solifuges (Arachnida, Solifugae) with special focus on functional analyses and phylogenetic interpretationsHISTOLOGIE UND ULTRASTRUKTUR DER SOLIFUGENVergleichende 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 NzurErlangung des akademischen Gradesdoctor rerum naturalium (Dr. rer. nat.)an der Mathematisch-Naturwissenschaftlichen Fakultät derErnst-Moritz-Arndt-Universität Greifswaldvorgelegt von Anja Elisabeth Klanngeboren am 28.November 1976 in BremenGreifswald, den 04.06.2009Prof. Dr. Klaus FesserDekan ........................................................................................................Prof. Dr. Dr. h.c. Gerd AlbertiErster Gutachter ..........................................................................................Prof. Dr. Romano DallaiZweiter Gutachter ........................................................................................15.09.2009Tag der Promotion ContentSummary .......................................................................................... 1Zusammenfassung ..........................................................................5Acknowledgments 91. Introduction ............................................................................. 121.1.
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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 04.06.2009Prof. 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 ............................................................. 177Content
3.12.1.2. Daesiidae .................................................................... 179
3.12.1.3. Eremobatidae ............................................................... 181
3.12.1.4. Galeodidae ........ 181
3.12.1.5. Karschiidae ....... 183
3.12.1.6. Solpugidae ........ 183
3.12.2. Discussion ......................................................................... 185
3.13. FOSSIL SOLIFUGE ........ 197
3.13.1. Results ............................................................................. 197
3.13.2. Discussion 198
3.14. MITOCHONDRIAL GENOME ...................................................... 200
3.14.1. Results 200
3.14.2. Discussion 200
4. Outlook ....................................................................................203
5. References ..............................................................................204
6. Appendix .................................................................................224
Publications .................................................................................225
Contributions to scientific meetings .........................................226
Eidesstattliche Erklärung ...........................................................228Summary
Solifuges (Solifugae, Arachnida) are an important element of the fauna especially
in arid and desert environments. Unfortunately, this animal group has only been
extremely poorly studied not only in terms of morphology, but also ecology, physiology,
systematics and phylogeny. The present study aimed to provide a detailed overview
of their anatomy and ultrastructure. Representatives of these peculiar animals
were investigated by means of (1) light and (2) electron microscopy in order to not
only predominantly gain new insights in their functional morphology, but also to
evaluate potential characters for their phylogeny.
Tarsal structures
Leg I-IV
Comparative scanning electron microscopic studies revealed differences in the
presence or absence of claws and arolia of leg I and the shape of the arolia of
the walking legs II-IV between the different families. The arolia of the walking
legs in the families Eremobatidae, Galeodidae and Solpugidae exhibit a family
specific groundpattern and not much variation. In contrast to these families, the
arolia in the families Ammotrechidae and Daesiidae show more variation in their
shape. Since the arolia of the walking legs of solifuges were never ultrastructurally
studied before, they could possibly provide new characters for their systematics
and phylogeny.
Suctorial organ
This peculiar organ is located on the tip of the distal tarsi of the pedipalp of
solifuges and is highly adhesive. Normally it is withdrawn and covered with cuticular
lips. The eversion process is probably triggered by increasing hemolymph pressure.
The suctorial organ itself is composed of numerous cuticular fibres with ramifying
apices fixed on a basal lattice-like, flexible plate. The fibers are covered with a
fragile epicuticle. Upon contraction of a muscle, the suctorial organ can rapidly be
withdrawn again.
Sensory structures and nervous system
Median eyes
The median eyes of solifuges are located on the ocular tubercle on the prosoma.
The prominent, spherical lens is underlain by a single cell layer, the lentigen layer.
The following retina consists exclusively of receptor cells. Apically the receptor
cells are provided with a surrounding microvilli border forming the rhabdomeres.
There are no additional pigment cells. The pigments are located in the receptor
cells.
1Summary
Nervous system
The prosomal nervous system of solifuges in general represents the groundpattern
of a typical arachnid brain. The supraoesophageal ganglion is composed of the
proto-, deuto- and the tritocerebrum and the suboesophageal ganglion consists of
several fused neuromeres. Two large tracheae enter the prosomal ganglion laterally
and ramify inside the brain. In contrast to scorpions and like uropygids there is only
one opisthosomal ganglion present in solifuges.
Malleoli
The present study mainly confirms former results on the malleoli being a highly
innervated sensory organ. The majority of the perikarya is concentrated in a
ganglion dorsally of each malleolus, which is located in the fourth pair of walking
legs. The numerous dendrites run through the malleolar stalk into the malleolar
fan. The outer dendritic segments end in the sensory groove. Very conspicuous are
numerous nuclei aggregated in the transition zone between the malleolar stalk and
malleolar fan.
Coxal gland
The coxal glands of the investigated solifuges consist of a (1) saccule, (2) mucous
segment, (3) a tubule and (4) an excretion porus. Apparently within the solifuges the
saccule is very fragile and associated with the mucous segment. The ultrastructure
of the mucous segment differs between the species: in contrast to Galeodes caspius
subfuscus the epithelial cells of the mucous segment of Nothopuga sp. contain
extremely large secretions. The epithelium of the tubule shortly before it merges
into the excretion porus consists of small cells and is folded. It does not exhibit
particular characteristics.
Alimentary System
The alimentary system of solifuges consists of the foregut (which comprises the
mouth, the pharynx and the oesophagus), the highly complex midgut and the hindgut.
Foregut and hindgut are provided with a cuticular intima. Solifuges masticate
their prey with their two-segmented chelicerae and ingest the particles through
a lattice at the tip of the rostrum (where the mouth is located) and transport
through the oesophagus and pharynx into the midgut. The resorption takes place
in the midgut, which consists of the midgut tube and numerous prosomal and
opisthosomal diverticula. The epithelium is composed of predominantly digestive
cells and secretory cells (which are less in numbers). Randomly, cells strongly
resembling endocrine cells in insect guts can be observed. The epithelium of the
midgut is basally interconnected with the intermediate tissue via so-called finger-
like processes. The epithelium of the hindgut is flat and overlain by a cuticle. The
main function of the hindgut is the storage and excretion of digested residuals
until defecation. The lateral branches similar to Malpighian tubules merge into
the midgut. They almost lack entirely cell inclusions, but possess high numbers of
mitochondria.
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