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15. Fossile Lebensgemeinschaften Im Lettenkeuper 15 Hans Hagdorn, Klaus-Peter Kelber Und Rainer Schoch

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15. Fossile Lebensgemeinschaften Im Lettenkeuper 15 Hans Hagdorn, Klaus-Peter Kelber Und Rainer Schoch 15. Fossile Lebensgemeinschaften im Lettenkeuper 15 Hans Hagdorn, Klaus-Peter Kelber und Rainer Schoch Abstract This chapter aims at reconstructing Lower Keuper palaeocommunities in their habitats and to elucidate interdependences of plants, invertebrates, and vertebrates. Four major groups of palaeocommunities are differentiated: (1) The terrestrial palaeocommunities are poorly known from transported fl oral and faunal elements into submersed depositional environments and rooted soil horizons. Clusters of insect eggs and nutritional traces on plants indicate animal-plant-interactions. Small pieces of charcoal give evidence of fi res. The terrestrial reptile record comprises small insectivores and larger carnivores with the rauisuchian Batrachotomus as the top predator, while herbivores are still unknown. The other groups of palaeocommunities depend on the salinity of the water bodies. (2) The fresh- water palaeocommunities are characterized by charophytes, by darwinulid ostracods, by bivalves (Unionites brevis/Unionites donaci- nus) and by amphibians. Here, a low diverse amphibian (Mastodonsaurus/Plagiosuchus/Gerrothorax) and a high diverse amphibian/ reptile palaeocommunity (Mastodonsaurus, Callistomordax, Kupferzellia, Trematolestes, choristoderes, and questionable juvenile tha- lattosaurs) can be discerned. (3) The three brackish water palaeocommunities are dominated by invertebrates: Myophoria/Bakevel- lia, Lingularia/Euestheria, “Anodonta” gregaria). (4) Among the nearly marine palaeocommunities are fi ve bivalve-dominated: Entolium discites, Pseudocorbula/Bakevellia subcostata, Hoernesia/Bakevellia subcostata with Placunopsis bioherms in Southern Baden-Würt- temberg, Costatoria goldfussi/Bakevellia subcostata, the latter two becoming more diverse towards the South, and the Central and North German Modiolus rhomboidalis/Costatoria goldfussi palaeocommunity documenting the northernmost expansion of the ma- rine Lower Keuper. One of the two marine vertebrate associations is dominated by diverse fi shes and reptiles plus the most saltwa- ter-tolerant amphibian, Plagiosternum, and another diverse palaeocommunity without amphibians. Stromatolites occur in three differ- ent horizons of the Lower Keuper and indicate hypo- or hypersalinar conditions. Non-quantitative examples of selected invertebrate dominated palaeocommunities from Northern Württemberg are ecologically characterised in terms of life habits and shell mineralogy. 1. Einführung haltes tolerierten, sowie an Wirbeltieren Amphibien und bestimmte Fische. Wenn die stratigraphische Zuordnung des Lettenkeu- Viele Ablagerungszyklen enden mit dickeren Karbo- pers lange Zeit zwischen dem marinen Muschelkalk und natbänken, den Leitbänken in der deskriptiven Lithostra- dem weitgehend terrestrischen Keuper geschwankt hat, tigraphie, die als marine Ingressionen kurzfris- so liegt das sicher auch an dem geographischen Stand- tige Überfl utungen des Beckens belegen (Kap. 13). Sie ort der Bearbeiter. Was für das räumliche Faziesmuster enthalten im südlichen Baden-Württemberg, wo sie fast der Sedimente gilt, trifft auch für die Verbreitung von Flo- durchgängig annähernd den ganzen Lettenkeuper auf- ren und Faunen zu. bauen, noch eine typische Muschelkalkfauna, wenn auch Unter der typischen „Lettenkohle“ wurde von den Au- stenohaline Elemente wie die Conodonten oder die Echi- toren des 19. Jahrhunderts ein komplexes Faziesmuster nodermen fehlen, die keine Abweichungen vom normalen aus tonigen Sandsteinen mit erosiv eingetieften Werksand- Salzgehalt tolerierten. Unter den Wirbeltieren dominieren stein-Rinnen, Wurzelhorizonten und dünnen Flözen toniger hier die Meeresreptilien und marine Fische. Nach Norden Kohlen verstanden. Dieser f l u v i a t i l e F a z i e s k o m - nimmt die Diversität dieser Fauna generell ab. So lag es p l e x (Kap. 13) ist besonders im zentralen und nördlichen nahe, dass die Bearbeiter der marin geprägten Lettenkeu- Beckenteil verbreitet. In diesen Sedimenten fi nden sich vor perfaunen Südwestdeutschlands (z.B. ALBERTI 1864) die allem Pfl anzen, aber auch Reste von Insekten, von Süß- Ähnlichkeiten mit dem Muschelkalk betonten und den Let- wassermuscheln und von Amphibien. tenkeuper zum Muschelkalk stellten, während die Faunen Nach Süden und innerhalb einzelner Ablagerungszyk- in Mittel- und Norddeutschland mehr den artenarmen Wir- len jeweils nach oben schließen sich dunkle, oft laminierte bellosenfaunen des Keupers entsprechen und folglich die Tonsteine mit karbonatischen Bänken an, die einen b r a - Zuordnung zum Keuper begründeten. ckisch-lakustrinen Ablagerungsraum Fossilien sind Ausnahmeerscheinungen im Kreislauf kennzeichnen (Kap. 13). An Wirbellosen fi nden sich da- des Werdens und Vergehens. Im Regelfall gehen Organis- rin vor allem Ostrakoden und Conchostraken sowie we- men in der Nahrungskette auf oder sie werden nach ihrem nige Arten von Muscheln, die Schwankungen des Salzge- Tod durch Zersetzung auf ihre anorganischen Grundbau- 359 PALAEODIVERSITY – LETTENKEUPER steine zurückgeführt. Die meisten Fossilien sind Reste von rem Vorkommen in bestimmten Gesteinsfazien ableiten. Meeresorganismen mit mineralisierten Skeletten. Weitaus Über trophische Beziehungen, also Beute/Räuber-Verhält- 15 seltener sind dagegen Fossilien von terrestrischen Lebens- nisse lässt sich angesichts des lückenhaften Fossilberichts formen – nicht zuletzt deshalb, weil terrestrische Sedimen- nur spekulieren. te weitaus häufi ger der Erosion zum Opfer fallen. Analysen der Vergesellschaftung von Wirbeltieren zei- Die Paläoökologie als Teildisziplin der Paläontologie gen sich ergiebiger als die ihrer stratigraphischen Reich- versucht nun als „Umweltwissenschaft“ die Lebensweise weiten (Kap. 4). Den zahlreichen unterschiedlichen Fossil- und Funktion ausgestorbener Organismen und deren Be- lagerstätten entsprechend lassen sich eine Reihe typischer, ziehungen zur unbelebten Natur zu verstehen. Aus dem mitunter häufi g wiederkehrender Assoziationen skizzieren. Fossilbefund ehemalige Lebensgemeinschaften zu rekon- Dabei bleibt zunächst offen, ob es sich um rein autochtho- struieren wird jedoch auf mehrfache Weise erschwert, ne, echte Lebensgemeinschaften handelt, oder ob sich da- denn (1) liegt stets nur der Teil einer ehemaligen Lebens- runter allochthone Elemente verbergen, und wenn, wie vie- gemeinschaft vor, der fossilisationsfähige Teile enthält le, d.h. ob es sich um Fossilvergesellschaftungen handelt. oder Spuren im Sediment hinterlassen hat, (2) durch se- Die Häufi gkeit und Vielgestaltigkeit der Bonebeds deutet ja lektiven Transport oder durch sedimentologische Prozesse bereits darauf hin, dass Transport und Umlagerung im Let- können einzelne Fossilien beigemischt oder entfernt sein, tenkeuper herausragend wichtige Faktoren waren. (3) der tatsächliche Bestand eines fossilführenden Hori- Fossilgemeinschaften und Vergesellschaftungen von zontes kann wegen ungünstiger Erhaltung, z.B. in Schil- Pfl anzenresten, Wirbellosen und von Wirbeltieren des Let- len im Festgestein, nur lückenhaft erfasst oder in seinen tenkeupers sind im Folgenden jeweils vom festen Land, quantitativen Verhältnissen nur geschätzt werden. Des- dann nach zunehmender Salinität vom Süßwasser über un- halb spricht man besser von f o s s i l e n L e b e n s g e - terschiedliche Brackwässer zum Meerwasser und schließ- m e i n s c h a f t e n ( f o s s i l e B i o z ö n o s e n ), und lich zum Salinarbereich angeordnet. Weil die triassischen auch dies nur im Falle von Wurzelhorizonten oder wenn Arten und Gattungen heute nicht mehr existieren, muss bei den Zweischalern Doppelklappigkeit, möglichst in Le- deren Habitatanspruch aus Fazies und Vergesellschaftung bendstellung vorliegt, bei den Wirbeltieren artikulierte Ske- der einzelnen Arten miteinander ermittelt werden. lette. Wenn Pfl anzenreste, Knochen oder Muschelklappen überwiegend isoliert und durch Transport eingeregelt sind, Abkürzungen handelt es sich um F o s s i l v e r g e s e l l s c h a f t u n - g e n ( T a p h o z ö n o s e n ), die auch gemischt sein Die Abbildungsoriginale sind in folgenden Sammungen hin- terlegt: SMNS – Staatliches Museum für Naturkunde Stuttgart; können, wenn einzelne Elemente aus anderen Lebensge- MHI – Muschelkalkmuseum Ingelfi ngen; GNO-, KREU-, OCH-, meinschaften hinzukommen. SCHL- Sammlung K.-P. KELBER, Würzburg. Weiteres Material wird Im Folgenden werden einige fossile Lebensgemein- in der Sammlung N. WANNENMACHER, Bisingen-Thanheim, aufbe- schaften bzw. Fossilvergesellschaftungen aufgeführt, die wahrt. sich in den Sedimenten des Lettenkeupers – mit Schwer- In den Faunenlisten der ausgewählten Fossilvergesellschaf- tungen sind folgende Abkürzungen verwendet: inf – Infauna; inft – punkt auf Nord-Württemberg – nachweisen lassen. Wo Infauna, tief grabend; sinf – Semiinfauna; mob – mobil; byss – möglich, wird ihr Kern, der aus den beiden jeweils domi- mit Byssus befestigt; zem – zementiert; susp – Suspensionsfi - nierenden Arten besteht, zur Benennung herangezogen. scher (Filtrierer); detr – Detritusfresser; weid – Weidegänger; Allerdings waren exakte quantitative Untersuchungen aus präd –Räuber und/oder Aasfresser (Prädator). den oben genannten Gründen nicht möglich. Wo die Schalenmineralogie bekannt ist, wird sie angegeben, Die Zusammensetzung der Lebensgemeinschaften und zwar wie folgt: A – Schale vollständig aragonitisch; KA – äußere Schalenschicht kalzitisch, innere aragonitisch; K – kalzi- des Lettenkeupers dürfte im Wesentlichen durch die Sali- tisch; P – phosphatisch. nität gesteuert worden sein. Ihre Differenzierung in der la-
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