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Microscopic Anatomy of Eukoenenia Spelaea (Palpigradi) — a Miniaturized Euchelicerate

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Microscopic Anatomy of Eukoenenia Spelaea (Palpigradi) — a Miniaturized Euchelicerate MICROSCOPIC ANATOMY OF EUKOENENIA SPELAEA (PALPIGRADI) — A MINIATURIZED EUCHELICERATE Sandra Franz-Guess Gröbenzell, Deutschland 2019 For my wife ii Diese Dissertation wurde angefertigt unter der Leitung von Herrn Prof. Dr. J. Matthias Starck im Bereich von Department Biologie II an der Ludwig‐Maximilians‐Universität München Erstgutachter: Prof. Dr. J. Matthias Starck Zweitgutachter: Prof. Dr. Roland Melzer Tag der Abgabe: 18.12.2018 Tag der mündlichen Prüfung: 01.03.2019 iii Erklärung Ich versichere hiermit an Eides statt, dass meine Dissertation selbständig und ohne unerlaubte Hilfsmittel angefertigt worden ist. Die vorliegende Dissertation wurde weder ganz, noch teilweise bei einer anderen Prüfungskommission vorgelegt. Ich habe noch zu keinem früheren Zeitpunkt versucht, eine Dissertation einzureichen oder an einer Doktorprüfung teilzunehmen. Gröbenzell, den 18.12.2018 Sandra Franz-Guess, M.Sc. iv List of additional publications Publication I Czaczkes, T. J.; Franz, S.; Witte, V.; Heinze, J. 2015. Perception of collective path use affects path selection in ants. Animal Behaviour 99: 15–24. Publication II Franz-Guess, S.; Klußmann-Fricke, B. J.; Wirkner, C. S.; Prendini, L.; Starck, J. M. 2016. Morphology of the tracheal system of camel spiders (Chelicerata: Solifugae) based on micro-CT and 3D-reconstruction in exemplar species from three families. Arthropod Structure & Development 45: 440–451. Publication III Franz-Guess, S.; & Starck, J. M. 2016. Histological and ultrastructural analysis of the respiratory tracheae of Galeodes granti (Chelicerata: Solifugae). Arthropod Structure & Development 45: 452–461. Publication IV Starck, J. M.; Neul, A.; Schmidt, V.; Kolb, T.; Franz-Guess, S.; Balcecean, D.; Pees, M. 2017. Morphology and morphometry of the lung in corn snakes (Pantherophis guttatus) infected with three different strains of ferlavirus. Journal of Comparative Pathology 156: 419–435. Publication V Starck, J. M.; Mehnert, L.; Biging, A.; Bjarsch, J.; Franz-Guess, S.; Kleeberger, D.; Hörnig, M. 2018. Morphological responses to feeding in ticks (Ixodes ricinus). Zoological Letters 4: 1–19. v Content Erklärung ................................................................................................................ iv List of additional publications................................................................................... v Summary ................................................................................................................ 1 1. Introduction ...................................................................................................... 2 1.1 Palpigradi and Miniaturization ............................................................. 2 1.2 Aims of the thesis ................................................................................ 6 2. Materials and Methods ..................................................................................... 7 2.1 Sampling and sourcing of specimens .................................................. 7 2.2 Scanning electron microscopy ............................................................. 9 2.3 Histology.............................................................................................. 9 2.4 Transmission electron microscopy .................................................... 10 2.5 Image processing .............................................................................. 11 2.6 Phylogenetic analysis ........................................................................ 11 2.7 Terminology ....................................................................................... 13 3. Results ........................................................................................................... 13 3.1 External morphology ......................................................................... 13 3.1.1 Prosoma dorsum ............................................................................... 14 3.1.2 Prosomal sternum ............................................................................. 14 3.1.3 Ventral plate ...................................................................................... 18 3.1.4 Rostrosoma ....................................................................................... 18 3.1.5 Sensory organs on the prosoma ....................................................... 20 3.1.6 Chelicerae ......................................................................................... 22 3.1.7 Pedipalps and legs ............................................................................ 23 3.1.8 Opisthosoma ..................................................................................... 24 3.1.9 Opisthosomal sclerites ...................................................................... 25 3.1.10 Genital segment ................................................................................ 26 vi 3.1.11 Flagellum ........................................................................................... 27 3.2 Internal morphology ........................................................................... 29 3.2.1 Epidermis .......................................................................................... 29 3.2.2 Cuticle ............................................................................................... 29 3.2.3 Endosternite ...................................................................................... 32 3.2.4 Muscle ultrastructure ......................................................................... 34 3.2.5 Musculature of the prosoma .............................................................. 35 3.2.5.1 Musculature of the extremities ................................................. 44 3.2.5.2 Musculature of the body wall ................................................... 45 3.2.5.3 Musculature of the endosternite ............................................... 48 3.2.6 Musculature of the opisthosoma ........................................................ 50 3.2.7 Nervous system ................................................................................. 53 3.2.8 Sensory organs ................................................................................. 59 3.2.8.1 Frontal organ ........................................................................... 60 3.2.8.2 Lateral organ ............................................................................ 64 3.2.8.3 Sensory setae .......................................................................... 65 3.2.8.4 Trichobothria ............................................................................ 68 3.2.9 Heart ................................................................................................. 71 3.2.10 Digestive tract .................................................................................... 73 3.2.10.1 Pharynx.................................................................................... 73 3.2.10.2 Esophagus ............................................................................... 73 3.2.10.3 Midgut ...................................................................................... 75 3.2.10.4 Rectal sac ................................................................................ 77 3.2.11 Excretory organ ................................................................................. 78 3.2.12 Reproductive organs ......................................................................... 82 3.2.12.1 Female ..................................................................................... 82 3.2.12.2 Male ......................................................................................... 86 3.2.13 Flagellum ........................................................................................... 90 vii 3.3 Phylogenetic analysis ........................................................................ 91 3.3.1 Phylogenetic position of Eukoenenia spelaea using Shultz's (2007a) original character matrix .................................................................... 91 3.3.2 Changes in character states and new characters in the data matrix . 95 3.3.3 Results for updated character matrix including data for Eukoenenia spelaea .............................................................................................. 99 3.3.4 Proposed apomorphic characters for Palpigradi and their sister group relationship ...................................................................................... 101 4. Discussion .................................................................................................... 102 4.1 Body tagmatisation .......................................................................... 102 4.1.1 Prosoma, dorsal aspect ................................................................... 103 4.1.2 Prosoma, ventral aspect .................................................................. 107 4.1.3 Chelicerae ....................................................................................... 109 4.1.4 Pedipalps and legs .......................................................................... 109 4.1.5 Opisthosoma ................................................................................... 110 4.2 Cuticle ............................................................................................. 114 4.2.1 Cutaneous respiration ..................................................................... 114 4.2.2 Surface structures ..........................................................................
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