Die Magenstrukturen Der Brachyura (Crustacea, Decapoda): Morphologie Und Phylogenetische Bedeutung

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Die Magenstrukturen Der Brachyura (Crustacea, Decapoda): Morphologie Und Phylogenetische Bedeutung Die Magenstrukturen der Brachyura (Crustacea, Decapoda): Morphologie und phylogenetische Bedeutung Dissertation zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) im Fach Biologie der Mathematisch-Naturwissenschaftlichen Fakultät I von Dipl.-Biol. Andreas Brösing, geb. am 28.06. 1968 in Dessau Präsident der Humboldt-Universität zu Berlin Prof. Dr. Jürgen Mlynek Dekan der Mathematisch-Naturwissenschaftlichen Fakultät I Prof. Dr. Michael Linscheid Gutachter: 1. Prof. Dr. Gerhard Scholtz 2. Prof. Dr. Hannelore Hoch 3. Prof. Dr. Johann-Wolfgang Wägele eingereicht: 20. September 2002 Tag der mündlichen Prüfung: 27. November 2002 Wissenschaft hat etwas Faszinierendes an sich. So eine geringfügige Investition an Fakten liefert so einen reichen Ertrag an Voraussagen. Mark Twain • Keywords: Brachyura, phylogeny, foregut-ossicles, fossil record • Schlagworte: Brachyura, Phylogenie, Magenossikel, Fossilbericht Abstract Within the Decapoda the taxon Brachyura is the species-richest taxon with up to 10000 species. The phylogeny of the Brachyura has been discussed based on morphological and molecular investigations since more than a century. The investigation of the foregut-ossicles and gastric-teeth of 66 brachyuran species, is a new approach to answer important phylogenetic questions. Using a specific staining pigment Alizarin Red S, six new described foregut-ossicles are added to an existing nomenclature. As a result of this method the presence of 41 foregut-ossicles is proposed for the ground pattern of the Brachyura. The cladistic analysis supports a monophyletic origin of the Brachyura including the Dromiidae and the Raninidae. The taxa Podotremata and Heterotremata, postulated as monophyletic by Guinot (1977, 1978), are not supported in the present study. The Dromiidae and Raninidae, which are placed within the Podotremata sensu Guinot, are closer related to the “higher crabs”. Based on the analysed foregut-characters, several molecular studies (Schubart et al. 2000a, 2000b), and the data of the fossil record, a monophyletic origin of the Thoracotremata sensu Guinot is suggested. The analysed basal position of the Dynomenidae and Dromiidae is in agreement with the brachyuran fossil record. A common ancestor for most of the “higher” brachyuran crabs is suggested for the period between the upper Cretaceous and the beginning of the Tertiary. Abstrakt Innerhalb der Decapoda stellt das Taxon Brachyura eines der artenreichsten Taxa mit bis zu 10000 Arten dar. Die Phylogenie der Brachyura wird aufgrund morphologischer und molekularer Untersuchungen seit mehr als ein Jahrhundert kontrovers diskutiert. Zur Klärung phylogenetischer Fragestellungen wurde mit den vergleichenden Untersuchungen der Magenossikel und der Magenzähne von 66 Taxa der Brachyura ein neuer phylogenetischer Ansatz gewählt. Mit Anwendung eines spezifischen Färbepigmentes, Alizarin-Rot S, konnten sechs neue Magenossikel einer bereits existierenden Nomenklatur hinzugefügt werden, so dass jetzt 41 Magenossikel für das Grundmuster der Brachyura angenommen werden können. Im Ergebnis der phylogenetischen Analyse wird ein monophyletisches Taxon Brachyura einschließlich der Taxa Dromiidae und Raninidae unterstützt. Die von Guinot (1977, 1978) als Monophyla postulierten Teiltaxa Podotremata und Heterotremata finden hier keine Unterstützung. Taxa wie Raninidae und Cymonomidae, welche aufgrund der coxalen Position ihrer Genitalporen dem Taxon Podotremata sensu Guinot zugeschrieben werden, weisen nach vorliegenden Daten eine nähere Verwandtschaft zu den „höheren“ Krabben auf. Eine Monophylie eines Taxons Thoracotremata sensu Guinot kann dagegen basierend auf den analysierten Magenstrukturen, aber auch auf der Grundlage molekularer Studien (Schubart et al. 2000a, 2000b) und dem vorliegenden Daten zum Fossilbericht angenommen werden. Eine basale Stellung der Dynomenidae und Dromiidae im Ergebnis der kladistischen Analyse lässt sich mit den Fossilfunden aus dem mittleren Jura in Übereinstimmung bringen. Des weiteren kann für die meisten Taxa der „höheren“ Krabben ein gemeinsamer Vorfahre für die obere Kreide bzw. bis zum Beginn des Tertiärs postuliert werden. Inhaltsverzeichnis 1 EINLEITUNG ............................................................................................................................... 1 1.1 STAND PHYLOGENETISCHER FORSCHUNGEN ZUM TAXON BRACHYURA................................ 1 1.2 KENNTNISSE ZUM FOSSILBERICHT DER BRACHYURA ............................................................. 3 1.3 STAND MAGENMORPHOLOGISCHER UNTERSUCHUNGEN ........................................................ 4 1.4 ZIELSETZUNG........................................................................................................................... 5 2 MATERIAL UND METHODEN................................................................................................. 6 2.1 ARTEN ..................................................................................................................................... 6 2.2 PRÄPARATION UND FÄRBETECHNIK........................................................................................ 7 2.3 RASTERELEKTRONENMIKROSKOPIE ........................................................................................ 8 2.4 DOKUMENTATION.................................................................................................................... 9 2.5 KLADISTISCHE ANALYSE ........................................................................................................ 9 3 ERGEBNISSE ............................................................................................................................. 10 3.1 ALLGEMEINE MORPHOLOGIE DES MAGENS .......................................................................... 10 3.2 ALLGEMEINE MORPHOLOGIE DER MAGENOSSIKEL UND MAGENZÄHNE.............................. 11 3.3 OSSIKEL-NOMENKLATUR...................................................................................................... 16 3.4 VERGLEICHENDE MORPHOLOGISCHE UNTERSUCHUNGEN DER MAGEN-OSSIKEL UND DER MAGENZÄHNE ................................................................................................................................... 18 3.4.1 Dromiidae de Haan, 1833.............................................................................................. 18 3.4.1.1 Magenossikel von Dromia spinirostris Miers, 1881................................................. 18 3.4.1.2 Magenossikel von Dromia wilsoni Fulton and Grant, 1902...................................... 21 3.4.1.3 Magenossikel von Dromia personata (Linnè, 1758)................................................. 21 3.4.1.4 Magenossikel von Lauridromia intermedia (Laurie, 1906) ...................................... 21 3.4.1.5 Magenossikel von Hypoconcha spinosissima Rathbun, 1933................................... 22 3.4.1.6 Magenzähne von Dromia wilsoni Fulton and Grant, 1902 ....................................... 23 3.4.1.7 Magenzähne von Dromia personata (Linnè, 1758) .................................................. 24 3.4.1.8 Magenzähne von Hypoconcha spinosissima Rathbun, 1933..................................... 26 3.4.2 Dynomenidae Ortmann, 1892 ....................................................................................... 27 3.4.2.1 Magenossikel von Dynomene filholi Bouvier, 1894 ................................................. 27 3.4.2.2 Magenossikel von Dynomene praedator Milne Edwards, 1879 ............................... 30 3.4.2.3 Magenossikel von Paradynomene tuberculata Sakai, 1963...................................... 30 3.4.2.4 Magenzähne von Paradynomene tuberculata Sakai, 1963 ....................................... 32 3.4.3 Homolidae de Haan, 1839 ............................................................................................. 33 3.4.3.1 Magenossikel von Dagnaudus petterdi (Grant, 1905) ............................................. 33 3.4.3.2 Magenossikel von Homola barbata (Fabricius, 1793).............................................. 36 3.4.3.3 Magenossikel von Homola ranunculus Guinot and Richer de Forges, 1995 ............ 36 3.4.3.4 Magenossikel von Paramola cuvieri (Risso, 1816) .................................................. 36 3.4.3.5 Magenzähne von Homola ranunculus Guinot and Richer de Forges, 1995.............. 38 3.4.4 Latreillidae de Haan, 1840 ............................................................................................ 39 3.4.4.1 Magenossikel von Latreillia valida de Haan, 1839................................................... 39 3.4.4.2 Magenossikel von Latreillia metanesa Williams, 1982 ............................................ 42 3.4.4.3 Magenzähne von Latreillia valida de Haan, 1839 .................................................... 43 3.4.5 Cymonomidae Bouvier, 1897........................................................................................ 45 3.4.5.1 Magenossikel von Cymonomus granulatus (Thompson, 1873) ................................ 45 3.4.5.2 Magenzähne von Cymonomus granulatus (Thompson, 1873).................................. 49 3.4.6 Raninidae de Haan, 1839............................................................................................... 50 3.4.6.1 Magenossikel von Ranina ranina (Linnaeus, 1758) ................................................. 50 3.4.6.2 Magenossikel von Raninoides hendersoni Chopra, 1933.......................................... 53 3.4.6.3 Magenossikel von Raninoides personatus
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