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Elasmobranch Egg Capsules Palaeoxyris, Fayolia and Vetacapsula As Subject of Palaeontological Research – an Annotated Bibliography

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Elasmobranch Egg Capsules Palaeoxyris, Fayolia and Vetacapsula As Subject of Palaeontological Research – an Annotated Bibliography See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/248399732 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula as subject of palaeontological research – an annotated bibliography Article · January 2008 CITATIONS READS 16 183 2 authors: Jan Fischer Ilja Kogan Urweltmuseum GEOSKOP Technische Universität Bergakademie Freiberg 95 PUBLICATIONS 304 CITATIONS 48 PUBLICATIONS 108 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Fishes from the Cretaceous of Saxony View project Fishes from the Cretaceous of Saxony View project All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Jan Fischer letting you access and read them immediately. Retrieved on: 10 November 2016 Freiberger Forschungshefte, C 528 psf (16) 75 – 91 Freiberg, 2008 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula as subject of palaeontological research – an annotated bibliography by Jan Fischer & Ilja Kogan, Freiberg with 6 figures FISCHER, J. & KOGAN, I. (2008): Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula as subject of palaeontological research – an annotated bibliography. – Paläontologie, Stratigraphie, Fazies (16), Freiberger Forschungshefte, C 528: 75–91; Freiberg. Keywords: Palaeoxyris, Fayolia, Vetacapsula, Scapellites, Spirangium, Crookallia, elasmobranch egg capsules, palaeoecology. Addresses: Dipl.-Geol. J. Fischer & Dipl.-Geol. I. Kogan, TU Bergakademie Freiberg, Geological Institute, Dept. of Palaeontology, Bernhard-von-Cotta street 2, 09599 Freiberg, Germany; [email protected] Contents: Abstract Zusammenfassung 1 Introduction 2 Discovery and confusion (1828–1887) 3 Insights and doubts (1888–1928) 4 Consensus and knowledge growth (1928–1985) 5 Palaeobiological approaches (1986–2008) 6 Conclusions Acknowledgements References Abstract This paper summarizes the 180 years history of research on the enigmatic fossil egg capsules Palaeoxyris, Vetacapsula and Fayolia. Originally described as plant remains, their interpretation was convincingly changed to egg capsules of ancient sharks by the end of the 19th century. The evolution of thought on them can generally be subdivided into four phases: starting from 1828 when the first specimens were discovered and described as plant remains; from 1888 their interpretation as egg capsules of sharks was introduced but caused controversy; between 1928 and 1985 this idea was widely accepted, the fossil record increased and probable producers were identified; finally, since 1986 the approach has turned from pure description and taxonomy to include multi- discipline analyses of their ecology and palaeobiogeography. The parataxonomic systematic of the capsules remains problematic. The most complete bibliography of this subject that has been compiled thus far concludes this contribution. Zusammenfassung Die Arbeit fasst die 180jährige Erforschungsgeschichte der rätselhaften fossilen Eikapseln Palaeoxyris, Vetacapsula und Fayolia zusammen. Von den ursprünglichen Beschreibungen als Pflanzenreste hat sich ihre Interpretation gegen Ende des 19. Jahrhunderts überzeugend hin zu Eikapseln von altertümlichen Haien gewandelt. Die Entwicklung der Ansichten über sie kann in vier Phasen unterteilt werden: 1828 beginnend, wurden erste Exemplare entdeckt und als Florenreste beschrieben; seit 1888 wurde ihre Deutung als Eikapseln von Haien vorgeschlagen, hat jedoch noch Kontroversen verursacht; zwischen 1928 und 1985 war diese Vorstellung weitgehend akzeptiert, die Anzahl der fossilen Nachweise nahm zu und es wurden mutmaßliche Erzeuger bestimmt; schließlich hat sich seit 1986 die Herangehensweise von einer reinen taxonomischen Beschreibung hin zu multidisziplinären Analysen ihrer Ökologie und Paläobiogeographie verändert. Die 75 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. _________________________________________________________________________________________________________________ parataxonomische Systematik der Kapseln bleibt problematisch. Die bisher vollständigste Bibliographie zu diesem Thema schließt diesen Beitrag ab. 1 Introduction The enigmatic fossil capsules Palaeoxyris BRONGNIART, 1828, Vetacapsula MACKIE, 1867 and Fayolia RENAULT & ZEILLER, 1884 have long been a source of conjecture and controversy. Originally described as plant remains, they experienced a number of assignments to numerous groups of plants throughout the 19th century. Their true identity as egg capsules of fossil sharks was not recognized until 1888, and it still took decades before this attribution was widely accepted. Since their discovery the knowledge of them grew continuously, especially regarding their structure, variety of shapes, geographical distribution, stratigraphical range and probable producers. In the last few years, the approach has turned from pure description and taxonomy to include multi-discipline analyses of their systematic, ecology and palaeobiogeography. The early workers used to summarize the preceding publications in order to be able to discuss different possible interpretations of the capsules. However, with the acceptance of the thesis of their elasmobranch origin, the necessity of such reviews has diminished. In the same time, the rising number of publications made it difficult to overview them all. Since most of the early papers are written in French, German, Swedish or Russian, they also pose a problem for the modern English-speaking scientific community. As a result, some basic works, published by influential palaeobotanists of the 19th and early 20th century, are largely unknown nowadays. Therefore, we intend to give an actual overview of the evolution of thought about these controversial fossils, summarizing the most important contributions and presenting the most complete bibliography of them that has been compiled so far. The paper is divided into four sections illustrating the main steps of knowledge progress during the last 180 years. 2 Discovery and confusion (1828-1887) In 1828, the founder of the modern palaeobotany, the French botanist A. BRONGNIART, described the first Buntsandstein (Middle Triassic, Anisian) flora from Sultz-les-Bains, Vosges, France. Among the different plant remains he noted two specimens of a spike-like structure covered with apparently regular, tile-like rhombic scales (Fig. 1). Interpreting them as an inflorescence similar to those of the recent angiosperm Xyris or the restiads (Fig. 4/1), BRONGNIART (1828a) assigned them to the new genus and species Palaeoxyris regularis. The interpretation as inflorescence and assignment to the xyrids or the restiads was also mentioned in BRONGNIART (1828b) and subsequently accepted by BRONN (1835–1837, 1837), UNGER (1845), GERMAR (1851), BRONN & ROEMER (1851–1852) and SCHMIDT (1855). _____________________________________________ Fig. 1: Illustration of the type specimens of Palaeoxyris regularis by BRONGNIART (1828a). Specimens measure circa 90 mm in length and 13 mm in width at their widest point. _____________________________________________ A second species, P. muensteri from the Keuper of Bamberg, Germany, was published by PRESL in STERNBERG (1838) who considered Palaeoxyris to be an appendage on a restiad. The first Palaeozoic specimen was described as Carpolites helicteroides (later placed in Palaeoxyris by KIDSTON, 1886) by MORRIS in PRESTWICH (1840) from the coal measures of Coalbrook Dale, England. Later, SCHIMPER & MOUGEOT (1844) mentioned noticeable size differences between P. regularis and P. muensteri. In 1850 a third species, P. carbonaria, was reported by SCHIMPER in STIEHLER (1850) from the Carboniferous shale clays of Wettin, Germany and compared with P. 76 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. _________________________________________________________________________________________________________________ regularis. He discussed similarities with the xyrids, but concluded most similarities with the bromeliads, especially for P. muensteri, a view which was subsequently shared by SCHENK (1864). ETTINGSHAUSEN in V. HAUER (1851) mentioned a new find from the Wealden of Deister, northern Germany, which he thought was close to the bromeliads. Therefore, ETTINGSHAUSEN (1852) erected the new taxon Palaeobromelia jugleri. It consisted of probably six spirally wound valves and as a result he believed it to differ from the scale-covered Palaeoxyris. Up to 12 capsules, which he considered as spike-like perianths of bromeliads (Fig. 4/2), were assembled in an umble-like formation in his reconstruction. An attempt to abscise Palaeoxyris from the xyrids was made by STIEHLER (1860) who proposed the new generic name Sporlederia. This name, however, was already preoccupied by a genus of mosses (BROWN, 1950). The first to express doubt about the plant origin of Palaeoxyris was BEER in 1856. In a short contribution he summed up the knowledge of Palaeoxyris and Palaeobromelia, removing the distinction between both genera and confirming the assignment of P. regularis to the bromeliads. P. muensteri and P. jugleri, however, were compared with an unlabeled specimen of unknown provenance but unquestionably animal origin in the collection of the imperial botanical museum of Vienna. It had been tentatively identified as the envelope of an egg by a colleague, K.A. FRAUENFELD. So BEER for the first time took Palaeoxyris into the animal realm. Unfortunately, BEER’s progressive view
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