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Did ammonoids possess opercula? Reassessment of phosphatised soft tissues in Glaphyrites from the Carboniferous of Uruguay

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Abstract

A reassessment of Glaphyrites material from the Late Carboniferous of Uruguay, famous for being the first ammonite described with the complete buccal mass in situ (also referred to Eoasianites earlier), indicates that the structures recorded as opercula or anaptychi, respectively, by earlier authors in several of the Glaphyrites specimens are rather to be interpreted as (1) dislodged fragments of the calcareous shell, originating from the aperture, and isolated by demineralization as well as fragmentation during taphonomy, and (2) remnants of demineralization, the fine lines suggesting that the rims of the ‘opercula’ are elongated “puckers” produced by shrinking of the inner layer of the calcareous shell during dissolution. An energy dispersive X-ray (EDX) analysis corroborated the impression that these fossils consist uniformly of a matrix of calcium phosphate (apatite), containing a less significant amount of iron and sulphur, among other elements. The Glaphyrites material is re-interpreted as possessing a lower and upper jaw with a general beak-like morphology that is comparable to modern nautilids and coleoids, a complex radula, but evidence for an operculum protecting the soft body is missing. Since the jaw elements are located in the anterior portion of the body chamber, but not directly at the aperture, and are furthermore of smaller size compared to the cross section of the aperture, we assume that a double function of jaws as opercula in Glaphyrites is unlikely.

Kurzfassung

Die erneute Untersuchung des Glaphyrites-Materials aus dem späten Karbon von Uruguay, berühmt als erste Ammonoideen bei denen die gesamte Buccalmasse in situ erhalten ist (früher z.T. auch Eoasianites zugeordnet), zeigt, daß die von früheren Autoren als Opercula bzw. Anaptychi beschriebenen Strukturen, die bei mehreren Exemplaren von Glaphyrites auftreten, eher als (1) dislozierte Fragmente der Kalkschale zu deuten sind, die aus dem Mündungsbereich stammen und während der Taphonomie durch Demineralisation und Fragmentierung isoliert und als (2) Demineralisationsreste vorliegen; die Runzeln, die an Opercula erinnern, sind durch den Schrumpfungsprozess der Innenseite der Kalkschale während deren Auflösung entstanden. EDX-Analysen bestätigen den makroskopischen Befund, dass diese Fossilien einheitlich aus einer Maxtrix von Calciumphosphat (Apatit) erhalten sind, welche Eisen und Schwefel enthält. Das Glaphyrites-Material wird hier dahingehend interpretiert, dass diese Ammonoideen Unter- und Oberkiefer von generell schnabelartiger Form besaßen, die grundsätzlich denen der modernen Nautiliden und Coleoideen ähneln, sowie eine komplexe Radula, aber kein Operculum. Da sich die Kieferelemente im vorderen Bereich der Wohnkammer finden, aber nicht direkt an der Mündung, und zu klein sind um den Windungsquerschnitt im Mündungsbereich abzudecken, halten wir eine Doppelfunktion als Kiefer und Deckel für Glaphyrites für unwahrscheinlich.

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Acknowledgments

H. Schulz (Tübingen) borrowed the specimens and kindly allowed ammonium chloride coating and sampling for SEM analysis; we also thank P. Havlik for additional information on the Glaphyrites specimens now under his care at the Tübingen collection. Technical support by M. Krogmann (Bremen) with respect to the macrophotographic works, as well as by P. Witte (Bremen) for SEM images, is highly appreciated. JL acknowledges funding by the German Research Foundation (DFG; project LE 1241/2-1). CK thanks the Swiss National Science Foundation for financial support (Project Number 132870). We thank both reviewers, H. Keupp (Berlin) and K. De Baets (Erlangen), for valuable comments that improved our manuscript.

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  1. Fachbereich Geowissenschaften, Universität Bremen, Klagenfurter Strasse, 28357, Bremen, Germany

    Jens Lehmann

  2. Paläontologisches Institut und Museum, Universität Zürich, Karl Schmid-Strasse 4, 8006, Zurich, Switzerland

    Christian Klug

  3. Anorganisch-chemisches Institut, Universität Zürich, Winterthurerstrasse 190, 5087, Zurich, Switzerland

    Ferdinand Wild

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Lehmann, J., Klug, C. & Wild, F. Did ammonoids possess opercula? Reassessment of phosphatised soft tissues in Glaphyrites from the Carboniferous of Uruguay. Paläontol Z 89, 63–77 (2015). https://doi.org/10.1007/s12542-013-0219-8

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