Elasmobranch Egg Capsules Palaeoxyris, Fayolia and Vetacapsula As Subject of Palaeontological Research

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

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 apparently had no impact on any subsequent research. SCHENK (1867) also pointed out that ETTINGSHAUSEN’s Palaeobromelia clearly belonged to Palaeoxyris and all differences were caused by preservation. He explained the appearance of the rhombic scales mentioned by preceding workers as a structure of six spirally wound leaves. If the fossil was flattened during compression, the windings on opposite sides would overlap, suggesting a scale-like rhombic pattern. Similar observations were made by QUENSTEDT (1867), who also mentioned a new find of Palaeoxyris (P. quenstedti SCHIMPER 1870– 1872) from the Keuper sandstone near Tübingen, Germany. He compared it to spirally wound leaves of cycads. In fact, SCHENK (1867) himself preferred to interpret Palaeoxyris as a spike-like cycad inflorescence. Furthermore, he pointed to the external similarities of Palaeoxyris and the egg cases of living plagiostoms (= chondrichthyian fishes). Despite these similarities, the differences were too significant to support an animal origin of Palaeoxyris.

______

Fig. 2: MACKIE’s type specimen of Vetacapsula cooperi (after CROOKALL, 1928b). Specimen measures 89 mm in length and 39 mm in width at its widest point. ______

Meanwhile, MACKIE (1867) erected Vetacapsula (“old capsule”) cooperi (Fig. 2), a new type of enigmatic flower-bud or seed-vessel (Fig. 4/4) consisting of two hemispheres with up to 26 ridges each from British coal measures. The first records of the capsules in the Palaeozoic of North America were presented by LESQUEREUX (1870) when he described three different specimens of Palaeoxyris from the Westphalian of Mazon Creek, Illinois. On this material LESQUEREUX was able to observe for the first time the remains of the flange (collarette) as outward protruding irregularly pointed triangular teeth or appendages on both sides of the body, together with close, very narrow striations on the whole surface, running in the direction of the windings. LESQUEREUX did not hazard any hypothesis on the systematic position of these enigmatic plant remains, a position he still argued ten years later (LESQUEREUX, 1880). The new generic name Spirangium (“spiral capsule”) was proposed for Palaeoxyris by SCHIMPER (1870–1872). Since the fossil remains had no connection with the living Xyris, he preferred the new name, which was only descriptive and did not suggest any uncertainty or false interpretation. This proposal was supported by SCHENK (1871) and ROMANOWSKI (1880), who described S. gilewi from the Triassic of East Turkestan on the basis of a 3-dimensional stone cast, interpreting it as a spirally wound spindle-like inflorescence. However, after reviewing the history of the problem, summarizing the synonyms, collecting localities, giving a diagnosis for the genus and all known British species, KIDSTON (1886) pointed out that Palaeoxyris still was the valid generic name. Although it would seemingly be appropriate with respect to the structure and the unclear systematic position of the capsule, Spirangium had to remain a taxonomic synonym because of the priority of the original name. Unlike his predecessors, NATHORST (1879) concluded from the aquatic flora and fauna that accompanied Palaeoxyris as well as from its general morphology resembling algal spores of recent charophytes, that it could represent some freshwater plant or charophycean algae. The aquatic habitat would also explain the long

77 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______stratigraphic range and limited variability of these fossils, which were uncharacteristic attributes of land plants. This view was critically discussed by SCHIMPER & SCHENK in ZITTEL (1890) A third form-type was published by RENAULT & ZEILLER (1884) from the Carboniferous coal mines of Commentry, France. It was named Fayolia in honor of the mines’ director FAYOL, and comprised two species F. dentata (Fig. 3) and F. grandis. The authors recognized important similarities with the problematic Palaeoxyris but distinguished them from the latter by some unique characteristics (its construction of only two valves in contrast to Palaeoxyris’ six, and explicit scars and spikes). Fayolia is arguably interpreted as seed-vessels similar to recent Medicago- or Hymenocarpus fructifications. At the same time, WEISS (1884) described a similar fossil from the Early Permian of Oberalben, Germany. This specimen, which WEISS regarded as an abnormal stem of a calamite (Fig. 4/3), was named Gyrocalamus but subsequently renamed Fayolia with respect to RENAULT & ZEILLER’s simultaneous (1884) work. One year later NEWBERRY (1885) announced two similar species from the Pennsylvanian of the United States, which he called Spiraxis without determining their true nature. WILLIAMS (1887) proposed for them the name Prospiraxis because Spiraxis was preoccupied by a gastropod genus. In the same year, WEISS (1887) reported another new species of Fayolia, F. sterzeliana, from the Carboniferous sandstone pits of Chemnitz-Borna, Germany. ______

Fig. 3: First illustration of Fayolia dentata by RENAULT & ZEILLER (1888). Specimen measures circa 120 mm in length and 25 mm in width at its widest point. ______

To summarize, in the first period between 1828 and 1887, the three capsule form-types Palaeoxyris BRONGNIART, 1828, Vetacapsula MACKIE, 1867 and Fayolia RENAULT & ZEILLER, 1884 were discovered and published and their close morphological similarity was recognized. BRONGNIART (1828a) described what he believed to be a new genus of fossil plants and his classification within the plant kingdom was followed by all subsequent authors. BRONGNIART’s “rhombic scales” covering the body of Palaeoxyris were definitely abandoned in 1867, when both QUENSTEDT and SCHENK recognized the taphonomic origin of this pattern. Hypotheses on the systematic placement varied across nearly every major group of vascular plants as well as the charophycean algae. Contradictory remarks by BEER (1856) and SCHENK (1867) were notable exceptions, but did not influence the whole discussion.

3 Insights and doubts (1888-1928)

New input came to the debate when SCHENK (1888) discussed the apparent differences between the French and German Fayolia species. He determined that the German specimens were clearly abnormal stems of calamites that should not even be assigned to Fayolia, whereas a specialist on recent plagiostoms needed to be consulted about the French fossils. For the second time, after his 1867 remark on Palaeoxyris, SCHENK suggested correlating these fossils with fishes. RENAULT & ZEILLER (1888), inspired by SCHENK’s proposal, conducted a detailed comparison between Fayolia specimens and recent heterodontid egg capsules of Cestracion [Heterodontus] philippi. They worked out the strong similarities between both types. Fayolia as well as the investigated recent egg capsules feature two helicoidal bands, which are accompanied by a more or less broad, membranous, lateral flange-like structure termed a collarette. Distinct narrow parallel striations are present on both bands and collarettes. Furthermore, the preparation of Fayolia capsules proved that their surface was not composed of epidermal cells as one would expect to find in any plant organ. Finally, they concluded that Fayolia as well as the morphological closely related Palaeoxyris definitely represent fish eggs similar to the egg capsules of Cestracion, Scyllium, rays and chimaers. Therefore, it can be said that the publication of RENAULT & ZEILLER (1888) marked the transfer of Fayolia and Palaeoxyris from the plant to the animal kingdom in the minds of many scientists. In addition, they suggested that Pleuracanthus [Xenacanthus] gaudryi from Commentry was the possible producer of F. dentata. A more detailed and precise argument for this view was provided in ZEILLER (1890), and repeated by BAUER et al. (1893). Previously, NATHORST (1889) had revised some Fayolia remains from Altenwald, Germany, which GOLDENBERG (1873) originally described as the arthropod Oniscina ornate, accepted RENAULT & ZEILLER’s (1888) view, and admitted that they were egg capsules of plagiostoms.

78 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

SAPORTA (1891) reviewed and evaluated preceding works up to 1884. At first he supported NATHORST’s (1879) interpretation, supposing even that the Palaeoxyris or, as he says, Spirangium capsules represented not some isolated vegetable organs (as no hints on their attachment to the plant had been found so far), but complete submerged aquatic plants. In a final supplement, however, he cited the newer works by SCHENK and RENAULT & ZEILLER from 1888, examining and agreeing with their arguments. Discussing among possible objections, that these fossils are found in freshwater deposits which are not common shark habitats, he noted that some recent sharks and rays are also known from freshwater areas like the Nicaragua lake. The fact that no skeletal shark fossils were found together with the capsules was explained by the very poor preservation potential of their cartilaginous skeleton. Nevertheless, he described a specimen from Pennsylvania sent to him by L. LESQUEREUX that showed two capsules of Spirangium (P.) intermedium associated with some kind of a stick, which was considered to be a neck spine of a shark, ray or chimaera. SEWARD (1894) described a new specimen of Fayolia (F. besti after CROOKALL, 1928) from the British lower coal measures sandstone of Barnard Castle. From comparisons with the hard and resistant horny cases of recent shark eggs he concluded that this character facilitated their preservation in sedimentary strata. In contrast, STAINIER (1894) interpreted the remains of Spiraxis interstrialis (Fayolia after CROOKALL, 1928) in the Late Devonian strata of Belgium as being in all probability the remains of algae. Also MACLOSKIE (1896) considered P. carbonaria the twisted fruits of tree-ferns. Although WHITE (1899) once again pointed to the priority of the generic appellation Palaeoxyris by BRONGNIART in contrast to SCHIMPER’s Spirangium, SAUVAGE (1905) described new remains of Spirangium from Lérida, Spain. He also distinguished between fossil egg capsules with one twist, which he regarded as eggs of the cestraciontidae, and those with a double row of crossing striae, that he assigned to holocephalians. SALFELD (1907) and KEILHACK (1908) noted again that all remains that were known came from deposits with no marine influence, which would make the interpretation as selachian egg capsules improbable. However, in a footnote SALFELD cited E. KOKEN’s comment about recent sharks swimming far upstream in rivers into continental realms in India. Crucial progress was made by MOYSEY (1910) when he defined the characteristic three-fold division of the Palaeoxyris capsule into pedicle, body and beak. MOYSEY also combined Spiraxis with Fayolia and considered these fossils as egg capsules of fishes. Moreover, he cited H. WOODWARD who suggested that these Coal measure fishes might have entered estuaries or rivers in order to deposit their egg capsules, and attached them to either floating or fixed vegetation. This would explain their common association with various plant remains and not with those of fishes. WOODWARD supposed that the superficial plant-like aspects of Palaeoxyris might have been a case of mimicry serving as a protective disguise for the embryo. In 1913, MOYSEY introduced a distinction between two generic forms (α and β) within Vetacapsula, on the basis of the existence of a medial ridge in some species. PRUVOST (1919) included a summary and discussion of the three genera in his explanations to the geologic map of northern France and described two species of Palaeoxyris as well as the new F. moyseyi. From a detailed evaluation of opposing arguments he confimed the earlier conclusion that these capsules must be understood as elasmobranchian eggs. Congruence of the stratigraphic range as well as of the geographical distribution would point to Pleuracanthus as the probable producer of Fayolia. Known as a freshwater shark, Pleuracanthus would not contradict Fayolia’s occurrence in freshwater strata. Palaeoxyris and Vetacapsula should be regarded as eggs of selachians; PRUVOST named Helodus as a possible producer of Palaeoxyris. These predominantly marine chimaeroid fishes would enter paralic basins, in which Carboniferous Palaeoxyris and Vetacapsula were found, and were also often associated with freshwater mollusks. Furthermore, a comparison of recent selachians would prove that these are not restricted to the sea, but spawn in rivers and even in freshwater lakes. An allied capsule type called Scapellites (from scapellus – skein), a rare type that seemed to be restricted to the Carboniferous of Belgium, was introduced by PRUVOST (1922) with Scapellites cottoni. SUTCLIFFE (1909) briefly reported new finds of P. prendeli, whereas new species of Palaeoxyris/Spirangium were described by DUN (1912) from the Triassic of Australia, by KAWASAKI (1925) from the Early Jurassic of Korea, and by CHABAKOV (1927) from the Carboniferous of the former Soviet Union. Whereas STERZEL (1918), PRUVOST (1919), FOURNIER (1925) and CHABAKOV (1927) considered Palaeoxyris, Fayolia and Vetacapsula as egg capsules of elasmobranchs, other authors (NINDEL, 1920; BEHM, 1924; SCHMIDT, 1928) still preferred a plant origin (Fig. 4/5). In his palaeobotanical overview GOTHAN (1914, 1921) listed Palaeoxyris and Fayolia as “problematica” with a still unclear systematic position.

In the second period, the general opinion on the origin of the capsules turned from different vegetable interpretations to egg capsules of fishes due mainly to the works of SCHENK (1888) and RENAULT & ZEILLER (1888) that were subsequently broadened by ZEILLER (1890), SAPORTA (1891) and PRUVOST (1919). Different potential producers, mostly selachians, were also discussed. Meanwhile, a small number of authors still argued for a plant origin. MOYSEY’s (1910) insights into the morphology as well as palaeoecology of the fossils mark a massive advancement in knowledge and influenced all subsequent work on fossil shark egg capsules.

79 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______

4 Consensus and knowledge growth (1928-1985)

Between 1928 and 1932 R. CROOKALL provided a complete and comprehensive overview of all species of Palaeoxyris, Fayolia and Vetacapsula known at the time (CROOKALL, 1928a, 1928b, 1930, 1932). Furthermore, at the same time he revised and erected several new species (P. warei, P. pringlei, P. edwardsi, P. duni, P. bohemica, F. cambriensis, F. eltringhami, F. besti, F. warei, F. interstrialis, V. kidstoni, V. moyseyi, and V. hemingwayi). In CROOKALL (1932) the nature and affinities of these fossils were discussed in detail. First he listed 12 different groups to which Palaeoxyris and related genera Fayolia and Vetacapsula had been referred or with which they had been compared. He demonstrated that all remains invariably occur in fresh- or brackish water deposits; the capsules were attached to plants by their pointed beaks, and never showed cellular issue, which precludes them from being plant remains. CROOKALL came to the conclusion that they were of animal origin and had to be accepted as egg capsules of elasmobranchs. Furthermore, CROOKALL cited D. WATSON’s remark about the shark family Hybodontidae, whose stratigraphical range covers exactly that of Palaeoxyris. With regard to Fayolia and Vetacapsula, other possible forms (Ichthyotomi, Petalodonttidae, Cochliodontidae, Cladontidae) were mentioned; their geological range, however, provided no assistance. PRUVOST (1930) presented Scapellites minor from Belgium, a second species of the rare capsule type Scapellites. SAMPELAYO (1946) reported the first record of Vetacapsula from the Carboniferous of Spain, which he erroneously described as P. marini. CHABAKOV (1949) presented the new species P. taurica from the Jurassic of the Crimea, Russia. More important was his subdivision of Vetacapsula, based on a characteristic already mentioned by MOYSEY (1913) and CROOKALL (1930): species without the noticeable medial ridge (V. kidstoni, V. hemingwayi and V. czernyshevi) were transferred into the new genus Crookallia. The first possible Mesozoic specimen of Palaeoxyris sp. from North America (Late Cretaceous of Kansas) published earlier by CROOKALL (1930), was redescribed by BROWN (1950). In the same paper he noted a remarkable resemblance between the Palaeoxyris capsule and those of living Chimaera species and also suggested that the noticeable “spine scars” in Fayolia might be breathing pores. The general acceptance of the fish egg origin was noted in MÄGDEFRAU’s (1953) textbook on palaeobotany, where the capsules were identified as selachian egg capsules, and comparable admissions were made by NINDEL (1955) and RANTZIEN (1956). Nevertheless, LANGFORD (1958), who summarized the Mazon Creek flora again regarded Palaeoxyris as a plant form such as a fructification. Among the several Palaeoxyris species that he listed was the new taxon P. ellipticus which in truth represents the first record of Fayolia from Mazon Creek biota. Some of LANGFORD’s Palaeoxyris specimens were attached to wood fragments by a tendril originating from the beak. DABER (1969), who published a new find of Palaeoxyris from Richtenberg, Germany, supposed that the ancient sharks from the shallow shelf areas of Northern England could have followed the large and slow river systems far into the continental realm for spawning. Whereas heterodontid eggs were similar to the Fayolia remains, Vetacapsula and Palaeoxyris resembled recent chimaeroid eggs. Based on a single specimen ZIDEK (1976) described P. lewisi from the Middle Pennsylvanian of Oklahoma. At the same time he made use of the opportunity to briefly discuss the nature and affinities of the Palaeozoic genera of egg capsules. He recognized that all the genera based on fossil egg capsules theoretically have to be regarded as form genera, containing generically unrelated species. Nonetheless, for practical purpose and future reference they still have to be described and labelled. Furthermore, ZIDEK undertook a detailed comparison of Palaeoxyris and modern screw-shaped heterodontid egg capsules. He demonstrated that both types share the characteristic three-fold division noted by other authors. In heterodontid eggs a spine-like structure comparable to the beak of Palaeoxyris is developed towards the attachment end but enveloped by broad collarettes, and therefore cannot be perceived immediately. Furthermore, the distal end of heterodontid egg capsules shows a cervix-like constriction similar to the body-pedicle transition in Palaeoxyris. So ZIDEK concluded that the difference between both types consists in the overall morphology rather than in the capsule construction. A possible relationship of heterodontids and hybodonts and the congruent geological range of the latter with Palaeoxyris as well as the typical occurrence in strata deposited in fresh- to brackish water environments made hybodonts appear the most probable producers of those capsules (Fig. 5). Interestingly, P. jugleri seemed to ZIDEK (1976) to largely differ from Palaeoxyris because of its whorled arrangement and he renamed it Spirangium jugleri to underline this exclusion. Largely influenced by this contribution, and following its arguments, MÜLLER (1978) gave an overview of the egg capsules of fossil chondrichthyans, with the main focus on Palaeoxyris. He investigated the morphological construction and important features of the capsules in detail and specified the most diagnostic characteristics. MCGHEE (1982) tried to correlate fossil egg capsules to possible producers on the basis of the size distribution of Palaeoxyris, Vetacapsula and Fayolia compared to those of living chondrichtyhans. From the almost total congruence between the allometric properties of chimaeroid and Palaeoxyris egg capsules he concluded that Palaeoxyris was possibly an ancient group of egg capsules with holocephalian affinity. However, Vetacapsula and Fayolia did not appear similar to any recent chondrichthyan egg cases. In the same year MCGHEE &

80 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

RICHARDSON (1982) described the first occurrence of Vetacapsula in North America, from the Westphalian of Mazon Creek, Illinois. New Palaeoxyris species were described by FRENTZEN (1932) with P. muelleri from Southern Germany and SZE (1954) with P. sino-coreanum from northwestern China. Further information on the fossil record of egg capsules were provided by KANIS (1956) for northwestern Spain, GALL (1971) for the Northern Vosges, France, BOERSMA (1973) for Hinzweiler, Germany, KAHLERT (1973) for the Flechtinger Höhenzug, Germany, BARTHEL et al. (1975) and SCHNEIDER et al. (1984) for the Saale Basin, Germany, and BAIRD et al. (1985a, 1985b) for the Braidwood freshwater fauna of Mazon Creek, Illinois. Synoptic overviews of the records from entire regions or countries were given by PRANTL (1933) for Czechoslovakia, WATERLOT (1934) for Saarland and Lorraine, and VAN DER HEIDE (1943) for the Netherlands. In connection with a new erected species, P. versabundus, from the northeastern Aral area, Russia, VIALOV (1984) tried to create a new parataxonomic nomenclature for fossil egg capsules. His scheme for the new taxa begins with the type Natisginia (fossil signs of birth); hierarchically downwards follow subtype Embriotheca, superclass Piscithecoidea (egg capsules of fishes), class Chondrychtytheca (egg capsules of chondrichthyans), subclass Elasmobranchythecia (egg capsules of elasmobranchs), order Selachithecida (selachian egg capsules), family Palaeoxyrisidae, genus Palaeoxyris, and finally the species. VIALOV’s (1984) nomenclature apparently has not been adopted by subsequent authors.

In the third period from 1928 to 1985, the idea of Palaeoxyris, Vetacapsula and Fayolia being the egg capsules of chondrichthyan fishes became widely accepted, the only exception being LANGFORD (1958). The principal subject of discussion during this period was now their assignment to the most probable producers. Generally the xenacanthids were mentioned as the producers of Fayolia whereas the chimaeroids and selachians and especially the hybodont sharks were considered the sources of the other types. Palaeoecological aspects also played an increasing role in the investigations. The overviews, observations and definitions by CROOKALL (1928–1932) were crucial during the following decades.

5 Palaeobiological approaches (1986-2008)

Inspired by the 1978 contribution of his predecessor A.H. MÜLLER, J.W. SCHNEIDER engaged in the taxonomical and palaeobiological problems of Fayolia and Palaeoxyris (SCHNEIDER, 1986, 1996; SCHNEIDER & REICHEL, 1989; RÖSSLER & SCHNEIDER, 1997; SCHNEIDER et al., 2003, 2005). Based on the size differentiation and frequency of two Palaeoxyris species and different hybodont shark remains from the Stephanian lacustrine Wettin strata of the Saale Basin, Germany, SCHNEIDER (1986) was able to assign both egg capsules to their putative producers (P. helicteroides to Lissodus lacustris and P. carbonaria to Limnoselache [Sphenacanthus SOLER-GIJÓN, 1997] vicinalis; this attribution was confirmed by GEBHARDT, 1988) (Fig. 5), which occurred in the same strata but in different sedimentary facies. The capsules were mainly found in fluviatile-palustrine deposits, whereas skeletal remains were restricted to pelagic lacustrine sediments. So it seemed plausible that both hybodont shark species used the shore area of the lake for spawning. Xenacanthids, who were the most probable producers of Fayolia (SCHNEIDER & REICHEL, 1989), migrated for spawning into fluviatile areas. An anadromous behavior, the migration from marine into freshwater for spawning, appeared to be improbable but could not be excluded. SCHNEIDER & REICHEL (1989) explained these intra-freshwater migrations by the advantages of higher oxygen concentration in turbulent water and biotic factors like food supply and protection from potential predators in specific nesting grounds, which were situated in marginal fluvial or lacustrine areas. The authors suggested a seasonal migration of oviparous female nonmarine sharks of the Carboniferous and Early Permian to particular spawning grounds more or less far from their normal habitat. This seasonal migration seemed to be a phylogenetically old pattern of behaviour, such as that found in modern oviparous sharks of the continental shelf. On the basis of the lithofacies and fossil association of egg capsule occurrences SCHNEIDER & REICHEL were able to visualize a scheme for the palaeogeographical positions of egg capsule finds and skeletal remains of sharks for the Mid European Permocarboniferous (Fig. 6). Furthermore, they redefined the diagnostic features of Fayolia and indicated that P. bohemica from the Westphalian of Nýřany, Czech Republic, in truth represents a Fayolia because of its cylindrical shape and the visible scar lines on its surface. Moreover, the distinct size difference in the teeth of the same taxon, detected in bed-by-bed sampling, appears to be an effect of changing biotope preference during ontogenesis, from the fluvial spawning grounds towards the lacustrine habitats typical for adult sharks (SCHNEIDER, 1996). The intraspecific variation of Fayolia specimens and thus the questionable value of Fayolia “species” was considered by RÖSSLER & SCHNEIDER (1997). Finally, SCHNEIDER et al. (2003, 2005) reported a Visean spawning ground in Saxony, Germany, with mass occurrences of Fayolia and sporadically Palaeoxyris, the oldest Palaeoxyris remains so far.

81 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______

Fig. 4: Diagram illustrating a few of the early theories about the plant nature of the fossil capsules: 1 – Palaeo- xyris as some kind of xyrid after BRONGNIART (1828); 2 – Palaeoxyris as some kind of bromeliad after ETTINGSHAUSEN (1852); 3 – Fayolia as abnormal calamite stem after WEISS (1884); 4 – Vetacapsula as seed vessel of Sigillaria after MACKIE (1867); 5 – Palaeoxyris as a cone-like seed-vessel after different authors (e.g. KIDSTON, 1886; BEHM, 1924).

In contrast SCHULTZE & SOLER-GIJÓN (2003, 2004) see in the presence of egg capsules in the European basins an indicator of marginal marine environment. They suppose that the reproduction strategy of these elasmobranchs indicates a explicit marine signal, as no recent oviparous shark is known to deposit its egg capsules in freshwater environment. Thus, elasmobranch egg capsules could even be used for detecting palaeosalinities. The first unequivocal Palaeoxyris species from the Mesozoic of North America was described by AXSMITH (2006) in a preliminary account, based on a small collection from the Late Triassic Chinle Formation in Arizona.

82 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

Fig. 5: Diagram illustrating the recent interpretation of Palaeoxyris as the screw-like egg capsules of hybodont sharks (e.g. CROOKALL, 1932; ZIDEK, 1976), especially of small Lissodus- or Lonchidion-like sharks (e.g. SCHNEIDER, 1986; FISCHER et al., in press). Here they are shown attached by tendrils to stems and branches of horsetails in specific nesting grounds in the shore areas of lakes and rivers.

Currently, a more comprehensive work has been submitted to formalize and expand on the preliminary study of these fossils (FISCHER et al., in press), which also contains an overview of recent elasmobranch oviparous behaviour. Additional Palaeozoic records of Palaeoxyris have been reported by GÓMEZ-PALLEROLA (1988) and SOLER- GIJÓN & POYATO-ARIZA (1995) from Lérida, Spain, by ZAJÍC (1988a, 1988b) from Czech Republic, by SCHULTZE et al. (1993) and SCHULTZE (1995) from Hamilton, Kansas, by AUE (2003) from Plötz, Germany, by HANNIBAL et al. (2003) from Knob Noster, Missouri, and most recently by ZESSIN (2008) from Majorca. Furthermore, Palaeoxyris was noted by GALL & GRAUVOGEL-STAMM (1993, 1999, 2005) as well as SELDEN &

83 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______

NUDDS (2007) in the northern Vosges, France, and by SROKA & RICHARDSON (1997) in the Mazon Creek biota. New Palaeozoic finds of Fayolia have been reported by POSCHMANN & SCHINDLER (1997) from Niedermoschel, Germany, and RONCHI & SANTI (2003) and SANTI (2007) from the Orobic Basin, Northern Italy. Synoptic overviews of the records from Northern France and Belgium were given by DERYCKE et al. (1995), from Czech Republic by ZAJÍC (2006, 2007, 2008) and ŠTAMBERG & ZAJÍC (2008), and from Saarland by POSCHMANN & SCHINDLER (1997) and HEIDTKE (2007). Additionally, POSCHMANN & SCHINDLER (1997) gave a tabular overview of all described species of Fayolia.

______

Fig. 6: Schematic diagram of the palaeobiogeo- graphical positions of egg capsule localities and skeletal remains of sharks in the freshwater basins of the Variscan orogen of the Carboniferous and Early Permian (after SCHNEIDER & REICHEL, 1989): 1 – Fayolia; 2 – Palaeoxyris; 3 – Orthacanthus; 4 – hybodonts; 5 – Xenacanthus/Bohemiacanthus; B – Borna-Ebersdorf (Erzgebirge Basin, Visean), fluvial sandstone; D – Dobis (northeastern Saale Basin, Stephanian C), shallow lacustrine mudstone; N – Nýřany (Central Bohemian Basin, Westphalian D), cannelcoal of an oxbow lake; O – Oberalben (Saar Basin, Lower Rotliegend), fluvial sandstone; P – Plötz (northeastern Saale Basin, Stephanian C), fluvial-palustrine roof shale of the coal seams; W – Weissig (Elbe Zone, Lower Rotliegend), fluvial sandstones in near shore lacustrine black siltstones; We – Wettin (northeastern Saale Basin, Stephanian C), black lacustrine bituminous mudstone. ______

FISCHER et al. (2007a, 2007b) and VOIGT et al. (2007) described two distinct types of capsules (Palaeoxyris sp. and a so far undetermined capsule type) from the fluvio-lacustrine deposits of the Triassic Madygen Formation in southwestern Kyrgyzstan. This discovery implies the presence of two different elasmobranch taxa, which used the freshwater environments of the Madygen Formation as spawning grounds. FISCHER et al. (2008a) showed that the Palaeoxyris remains from the Late Triassic Chinle Formation of Arizona, with their combination of unusually well preserved collarettes and striations, appear surprisingly similar to the design of modern heterodontid egg capsules, resembling them in both capsule construction and overall shape. FISCHER et al. (2008b) mentioned that a total of 31 species of Palaeoxyris have been described so far, including specimens, which were obviously wrongly assigned to Palaeoxyris, and a further 11 records not classified to species level. The stratigraphical record extends about 250 million years from the Early Carboniferous (Visean) to the Late Cretaceous, with a notable gap throughout the Permian (mentioned previously by SCHNEIDER & REICHEL, 1989). Moreover, FISCHER et al. (2008b) detected a morphological criterion to distinguish pre- and post-Permian examples of Palaeoxyris consisting of the rib pattern on the pedicle. The pedicle of pre-Permian specimens is invariably traversed by spiral ribbing structures, forming a rhomboidal pattern over its whole length. In all known Mesozoic specimens, in contrast, the pedicle is marked by nearly longitudinally arranged ribs, which gradually taper towards its end. The reason for this change is still unknown and obscured by the Permian gap.

In the fourth period the attribution of the egg capsules to their probable producers was substantiated by new finds and methods. Beginning with SCHNEIDER & REICHEL (1989), facies analysis of the localities and detailed studies of the skeletal remains provided insights into the spawning behaviour of the ancient sharks and habitat preferences during ontogenesis. Two opposing assumptions were proposed regarding the nonmarine palaeoecology of these shark taxa and the correct interpretation of the egg capsules: permanent freshwater fishes with intra-freshwater spawning behaviour (SCHNEIDER & REICHEL, 1989) versus euryhaline anadromous fishes (SCHULTZE & SOLER-GIJÓN, 2004).

84 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

6 Conclusions

Known for 180 years, the interpretation of Palaeoxyris, Fayolia and Vetacapsula has fundamentally changed from enigmatic plant remains to the egg capsules of fossil sharks. Among the early publications assigning them to plants, BRONGNIART (1828a), ETTINGSHAUSEN (1852), QUENSTEDT (1867), SCHENK (1867) and SCHIMPER (1870–72) were of great influence. RENAULT & ZEILLER (1888) marked the turning point in the interpretation; further milestones were the works by MOYSEY (1910), PRUVOST (1919), CROOKALL (1928-1932), ZIDEK (1976) and SCHNEIDER & REICHEL (1989). In contrast, contributions written in languages other than English, French or German (e.g. CHABAKOV, 1949 or VIALOV, 1984) received little attention because of the linguistic barrier, independent of their true value, necessity or accuracy. It is thus very probable, unfortunately, that a number of interesting works still remain unknown to most researchers in Europe and America. The principal arguments pro and contra regarding the assignment of the capsules to fossil sharks can be summarized as follows: in favour of a vegetable interpretation, there were (1) an external similarity to inflorescences or fructifications of some recent plants, (2) their frequent occurrence together with different plant remains, (3) the fact that they mostly were found in shallow fresh water deposits that were not believed to be habitats of sharks, and (4) their rare associations with skeletal remains of sharks. On the other hand, (1) the structure of the capsules is seldom carbonized and never shows a cellular pattern as plant organs do, (2) detailed comparisons with recent chimaeroid and especially elasmobranch egg capsules show considerable similarities in both overall shape and capsule construction; and finally, (3) their stratigraphical range does not agree with that of the compared groups of plants, but is concordant with the development of chondrichthyans. Furthermore, on the basis of the comparison with the spawning behaviour of recent marine oviparous sharks and facies analyses of the fossil bearing strata, it is probable that ancient sharks might have migrated to specific freshwater spawning grounds far from their normal habitat, either marine or non-marine. As a result, the attribution of these capsules to fossil sharks is very probable but cannot be claimed as absolute because no specimen showing definite evidence (like a capsule containing a fish embryo or an egg capsule within a shark body fossil) has yet been found. However, a well-established egg-producer relationship exists for Palaeoxyris and hybodonts, with two documented cases of direct assignment of a specific Palaeoxyris species to its putative source (SCHNEIDER, 1986; FISCHER et al., in press), whereas the attribution of Fayolia to xenacanthids is regarded to be most plausible. In contrast, the producers of Vetacapsula and Scapellites are still unknown. Anyhow, the classification of these fossils is parataxonomic, consisting of form groups because of their ambiguous orthotaxonomic position. Previously PRUVOST (1919, 1930) remarked that all generic and specific names used for the chondrichthyan egg capsules have to remain provisonal until their attribution to producers is clear. Later workers like ZIDEK (1976) and MÜLLER (1978) pointed out that generic and specific categories of the egg capsules are not necessarily congruent with those of their producers. RÖSSLER & SCHNEIDER (1997) noted that in case of Fayolia, the diagnoses of the species are often doubtful. Differences might be caused by intraspecific variation (as it could be seen on a single find locality) rather than by distinct specific characteristics. Nevertheless, they are typically treated under the International Rules of Zoological Nomenclature (IRZN) for practical use and future reference (ZIDEK, 1976; RÖSSLER & SCHNEIDER, 1997). With this contribution, a more accessible basis is given for further works. The genera Palaeoxyris, Fayolia, Vetacapsula and Scapellites are in need of revision. Research is in progress by the Freiberg working group. New finds and modern methods of palaeobiology, sedimentology and geochemistry should provide a better understanding of the palaeoecology of the capsules and their producers.

Acknowledgements

We are thankful to our teacher J.W. SCHNEIDER (Freiberg University), whose works inspired us to this contribution and who supported it with useful comments. We are indebted to S.R. ASH (University of New Mexico, Albuquerque) for English editing and critical and helpful review that greatly improved the manuscript. Thanks also go to B.J. AXSMITH (University of South Alabama), S. YANBIN (Institute of Geology and Palaeontology, Nanjing), and N. UDOVICHENKO (University of Lugansk, Ukraine) for providing relevant literature. We want to thank O. ELICKI and other colleagues from our Department, who provided considerable progress of our work through controversial discussions. The reconstructions of the two opposed interpretation models were drawn by F. SPINDLER (Freiberg University) under supervision by the authors. The German Research Foundation (DFG) supports JF’s work with the grant SCHN 408/14-1.

85 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______

References

AUE, J. (2003): Zwei fossile Haifisch-Eikapseln auf einer Tonschieferplatte des Oberkarbon. – Arbeitskreis Paläontologie Hannover, 31: 95–98. AXSMITH, B.J. (2006): The first Mesozoic record of the enigmatic fossil Palaeoxyris from North America; Chinle Formation, Petrified Forest National Park. – In: PARKER, W.G., ASH, S.R. & IRMIS, R.B. (eds.): A Century of Research at Petrified Forest National Park: Geology and Paleontology. Museum of Northern Arizona Bulletin, 62: 1–2. BAIRD, G.C., SHABICA, C.W., ANDERSON, J.L. & RICHARDSON, J.E.S. (1985a): Biota of the Pennsylvanian muddy coast: habitats within the Mazonian Delta Complex, Northeast Illinois. – Journal of Paleontology, 59 (2): 253–281. BAIRD, G.C., SROKA, S.D., SHABICA, C.W. & BEARD, T.L. (1985b): Mazon Creek-type fossil assemblage in the U.S. midcontinent Pennsylvanian: their recurrent character and palaeoenvironmental significance. – Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 311 (1148): 87– 99. BARTHEL, M., MÜTZE, K. & SIMON, R. (1975): Neue Funde fossiler Pflanzen aus dem Saale-Trog. – Wissenschaftliche Zeitschrift der Humboldt-Universität zu Berlin. Math.-Nat. Reihe, 24 (4): 475–485. BAUER, M., DAMES, W. & LIEBISCH, T. (1893): Pflanzen. – Neues Jahrbuch für Mineralogie, Geologie und Palaeontologie, Jahrgang 1893 (I. Band): 203–204. BEER, J.G. (1856): Einiges über Bromeliaceen. – Bonplandia - Zeitschrift für die gesammte Botanik. IV. Jahrgang (No. 24): 382–384. BEHM, H.W. (1924): Vor der Sintflut. Ein Bildatlas aus der Vorzeit der Welt. – Franckh'sche Verlagshandlung, Stuttgart: 43. BOERSMA, M. (1973): Die Stefanflora von Hinzweiler. – Courier Forschungsinstitut Senckenberg, 1: 10–12. BRONGNIART, A. (1828a): Essai d'une Flore du grès bigarré. – Annales des sciences naturelles, 15: 435–460, Pl. XX. BRONGNIART, A. (1828b): Prodrome d'une histoire des végétaux fossiles. – F. G. Levrault, Paris: 223 pp. BRONN, H.G. (1837): Allgemeine Encyklopädie der Wissenschaften und Künste. Dritte Section O–Z. – In: ERSCH, J.G. & GRUBER, J.G. (eds.). Brockhaus, Leipzig: 500 pp. BRONN, H.G. (1835–1837): Lethaea Geognostica. Erster Band. – E. Schweizerbart's Verlagshandlung, Stuttgart: 150–151. BRONN, H.G. & ROEMER, F. (1851–1852): Lethaea Geognostica. Zweiter Band. – E. Schweizerbart'sche Verlagshandlung, Stuttgart: 412 pp. BROWN, R.W. (1950): Cretaceous fish egg capsule from Kansas. – Journal of Paleontology, 24 (5): 594–600. CHABAKOV, A.W. (1927): O nachodke w kamennougolnych otloshenijach SSSR jajzewych kapsul elasmobranchij Palaeoxyris i Vetacapsula. [About the find of elasmobranch egg capsules Palaeoxyris and Vetacapsula from Carboniferous deposits of the USSR]. – Westnik geologitscheskogo komiteta, 1: 33–34 [in Russian]. CHABAKOV, A.W. (1949): O dwuch nowych formach problematitschnych iskopayemych Crookallia is karbona Donetzkogo basseyna i Palaeoxyris is jury Krima. [About two new problematic fossils Crookallia from the Carboniferous of Donezk Basin and Palaeoxyris from the Jurassic of Crimea]. – Annual of the All-Union Paleontological Society, 13: 83–87, Pl. IX [in Russian]. CROOKALL, R. (1928a): The genus Fayolia. – The Naturalist: 325–332 & 361–367, Pl. VIII–IX. CROOKALL, R. (1928b): Palaeozoic species of Vetacapsula and Palaeoxyris. – Summary of Progress of the Geological Survey of Great Britain and the Museum of Practical Geology for 1927, part II: 87–108, Pl. I–II. CROOKALL, R. (1930): Further morphological studies in Palaeoxyris, etc. – Summary of Progress of the Geological Survey of Great Britain and the Museum of Practical Geology for 1929, part III: 8–36, Pl. II–V. CROOKALL, R. (1932): The nature and affinities of Palaeoxyris, etc. – Summary of Progress of the Geological Survey of Great Britain and the Museum of Practical Geology for 1931, part II: 122–140, Pl. II. DABER, R. (1969): Paläobotanische Hinweise auf eine paralisch beeinflußte Oberkarbon-Senke im tieferen Untergrund Nordostdeutschlands, II (Teil 1). – Geologie, 18 (3): 253–297. DERYCKE, C., CLOUTIER, R. & CANDILIER, A.-M. (1995): Palaeozoic vertebrates of northern France and Belgium: Part II – Chondrichthyes, Acanthodii, Actinopterygii (uppermost Silurian to Carboniferous). – Geobios, 19: 343–350. DUN, W.S. (1912): Note on the occurrence of the genus Spirangium in the Hawkesbury Series of New South Wales. – Journal and Proceedings of the Royal Society of New South Wales XLVI: 205–206, Pl. XIV. ETTINGSHAUSEN, C.v. (1852): Ueber Palaeobromelia, ein neues fossiles Pflanzengeschlecht. – Abhandlungen der kaiserlich-königlichen geologischen Reichsanstalt I (3. Abth.): 1–10, Pl. I–II.

86 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

FISCHER, J., AXSMITH, B.J. & ASH, S.R. (in press): First unequivocal record of the hybodont shark egg capsule Palaeoxyris in the Mesozoic of North America. – Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen. FISCHER, J., AXSMITH, B.J. & ASH, S.R. (2008a): Resemblance of Palaeoxyris and modern heterodontid egg capsules. – Jahrestagung der Paläontologischen Gesellschaft, GeoZentrum Nordbayern, 8.–10. September 2008, Erlangen, Erlanger Geologische Abhandlungen, Institut für Geologie der Universität Erlangen- Nürnberg, Sonderband, 6: 85. FISCHER, J., SCHNEIDER, J.W. & BUCHWITZ, M. (2008b): Pedicle ribbing structure of the egg capsule Palaeoxyris reflects a Palaeozoic to Mesozoic change-over in its chondrichthyan producers. – Jahrestagung der Paläontologischen Gesellschaft, GeoZentrum Nordbayern, 8.–10. September 2008, Erlangen, Erlanger Geologische Abhandlungen, Institut für Geologie der Universität Erlangen-Nürnberg, Sonderband, 6: 85–86. FISCHER, J., VOIGT, S. & BUCHWITZ, M. (2007a): Two distinctive elasmobranch egg capsules from freshwater deposits of the Madygen Formation (Middle to Upper Triassic, Southwestern Kyrgyzstan, Central Asia). – 77. Jahrestagung der Paläontologischen Gesellschaft, Freiberg, 17.–19. September 2007, abstracts; Wissenschaftliche Mitteilungen des Institutes für Geologie, 36: 39. FISCHER, J., VOIGT, S. & BUCHWITZ, M. (2007b): First elasmobranch egg capsules from freshwater lake deposits of the Madygen Formation (Middle to Late Triassic, Kyrgyzstan, Central Asia). – Freiberger Forschungshefte, C 524: 41–46. FOURNIER, E. (1925): Les Fayolia sont bien des coques d'ceufs d'Elasmobranches. – Compute rendu sommaire des séances de la Société Géologique de France: publication bimensuelle, 9: 128–130. FRENTZEN, K. (1932): Beiträge zur Kenntnis der fossilen Flora des südwestlichen Deutschland. IX. Revision der Rhätflora Schwabens. – Jahresberichte und Mitteilungen des Oberrheinischen geologischen Vereines - Neue Folge, Band XXI: 75–94. GALL, J.-C. (1971): Faunes et paysages du Grès à Voltzia du Nord des Vosges. Essai paléoécologique sur le Buntsandstein supérieur. – Mémories du Service de la Carte géologique d'Alsace et de Lorraine 34. Université Louis Pasteur de Strasbourg, Strasbourg: 318 pp. GALL, J.-C. & GRAUVOGEL-STAMM, L. (1993): Paleoecology of terrestrial ecosystems from the Buntsandstein (Lower Triasic) of Eastern France. – New Mexico Museum of Natural History and Science Bulletin, 3: 141– 145. GALL, J.-C. & GRAUVOGEL-STAMM, L. (1999): Paläoökologie des Oberen Buntsandsteins am Westrand des Germanischen Beckens: Der Voltziensandstein im nordöstlichen Frankreich als deltaische Bildung. – In: HAUSCHKE, N. & WILDE, V. (eds.): Trias - Eine ganz andere Welt. Verlag Dr. Friedrich Pfeil, München: 283– 299. GALL, J.-C. & GRAUVOGEL-STAMM, L. (2005): The early Middle Triassic 'Grès à Voltzia' Formation of eastern France: a model of environmental refugium. – Comptes Rendus Palevol, 4 (6–7): 637–652. GEBHARDT, U. (1988): Taxonomie und Palökologie von Lissodus lacustris n. sp. (Hybodontoidea) aus dem Stefan C (Oberkarbon) der Saalesenke. – Freiberger Forschungshefte, C 419: 38–44. GERMAR, E.F. (1851): Die Versteinerungen des Steinkohlengebirges von Wettin und Löbejün im Saalekreise. – 7. Heft. c. A. Schwetschke und Sohn, Halle: 95–96, Pl. XXXIII. GOLDENBERG, F. (1873): Die fossilen Thiere aus der Steinkohlenformation von Saarbrücken. – Fauna Saraepontana Fossilis Heft 1. Verlag von Chr. Möllinger, Saarbrücken: 26 pp. GÓMEZ -PALLEROLA, J.E. (1988): Nota sobre los peces elasmobranquios de las calizas litográficas del Cretácico Inferior del Montsec (Lérida). – Boletin Geológico y Minero, 99 (5): 748–756. GOTHAN, W. (1914): Die unter-liassische ("rhätische") Flora der Umgegend von Nürnberg. – Abhandlungen der Naturhistorischen Gesellschaft zu Nürnberg XIX. Band (IV): 162. GOTHAN, W. (1921): Lehrbuch der Paläobotanik. – zweite Auflage, umgearbeitet. Verlag von Gebrüder Borntraeger, Berlin: 537. HANNIBAL, J.T., KEIPER, J.B., LEMAY, S. & MCKENZIE, S. (2003): Knob Noster; a new Upper Carboniferous (Pennsylvanian) lagerstätte in Missouri containing millipeds (Diplopoda), insects, crustaceans, vertebrates, plants and other terrestrial and brackish water organisms. – Abstracts with Programs - Geological Society of America, 35 (2): 60. HEIDTKE, U.H.J. (2007): Räuber in Flüssen und Seen – Haie im Süßwasser. – In: SCHINDLER, T. & HEIDTKE, U.H.J. (eds.): Kohlesümpfe, Seen und Halbwüsten. Dokumente einer rund 300 Millionen Jahre alten Lebewelt zwischen Saarbrücken und Mainz. Pollichia Sonderveröffentlichung, 10: 206–229. KAHLERT, E. (1973): Die biostratigraphische Einstufung der "Süplinger Schichten" (Flechtinger Höhenzug). – Zeitschrift für geologische Wissenschaften, 1 (7): 851–859. KANIS, J. (1956): Geology of the Eastern Zone of the Sierra del Brezo (Palencia-Spain). – Leidse Geologische Mededelingen, 21 (2): 377–445. KAWASAKI, S. (1925): Some older Mesozoic plants in Korea. – Bulletin on the Geological Survey of Chosen (Korea) IV (1): 57–58, Pl. XLVII.

87 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______

KEILHACK, K. (1908): 2472. Salfeld. – Geologisches Centralblatt - Anzeiger für Geologie, Petrographie, Palaeontologie und verwandte Wissenschaften, Band X: 733–734. KIDSTON, R. (1886): On the Species of the Genus Palaeoxyris, BRONGNIART, occurring in British Carboniferous Rocks. – Proceedings of the Royal Physical Society of Edinburgh, 9: 54–64, Pl. I. LANGFORD, G. (1958): The Wilmington coal flora from a Pennsylvanian deposit in Will County, Illinois. – Esconi Associates, Downers Grove: 295–302. LESQUEREUX, L. (1870): Organs of uncertain affinity. – Geological Survey of Illinois. IV, Springfield: 464–467, Pl. XXVII. LESQUEREUX, L. (1880): Description of the coal flora of the Carboniferous Formation in Pennsylvania and throughout the United States. – Second Geological Survey of Pennsylvania: Report in Progress Vol. II. Geological Survey, Harrisburg: 518–521, Pl. LXXV. MACKIE, S.J. (1867): Undescribed Vegetable Fossil. – The Geological and Natural History Repertory, Vol. i (1865–67): 79–80. MACLOSKIE, G. (1896): Further Observations on Antidromy. – Bulletin of the Torrey Botanical Club, 23 (10): 420–423. MÄGDEFRAU, K. (1953): Paläobiologie der Pflanzen. – Gustav Fischer Verlag, Jena: 483 pp. MCGHEE JR., G R. (1982): The problematic Palaeoxyris, Vetacapsula, and Fayolia: a morphological comparison with recent chondrichthyan egg cases. – North American Paleontological Convention, Proceedings, 2: 365– 369. MCGHEE JR., G.R. & RICHARDSON JR., E.S. (1982): First occurrence of the problematic fossil Vetacapsula in North America. – Journal of Paleontology, 56 (5): 1295–1296. MOYSEY, L. (1910): On Palaeoxyris and other allied fossils from Derbyshire and Nottinghamshire Coalfield. – The quarterly journal of the Geological Society of London, 66: 329–345, Pl. XXIV–XXVII. MOYSEY, L. (1913): Some further Notes on Palaeoxyris and other allied Fossils, with Special Reference to some New Features found in Vetacapsula. – British Association for the Advancement of Science: 492–493. MÜLLER, A.H. (1978): Über Palaeoxyris und andere Eikapseln fossiler Knorpelfische (Chondrichthyes). – Freiberger Forschungshefte, C 342: 7–28. NATHORST, A.G. (1879): Om Spirangium och dess förekomst. – I. Skånes. Kolförande Bildningar. Svergies Geologiska Undersökning Ser. C, 36: 1–13, Pl. VI–VII. NATHORST, A.G. (1889): Ueber Goldenberg's Onisima ornata. – Zeitschrift der Deutschen geologischen Gesellschaft, 41: 545–547. NEWBERRY, J.S. (1885): Description of some peculiar screw-like Fossils from the Chemung Rocks. – Annales of the New York Academy of Sciences, 3 (7): 217–220, Pl. XVIII. NINDEL, F. (1920): Ein Beitrag zur Fayolia sterzeliana (Ch. E. Weiß) aus dem Ober-Kulm von Chemnitz-Borna. – Zwanzigster Bericht der Naturwissenschaftlichen Gesellschaft zu Chemnitz: 49–54. NINDEL, F. (1955): Die tierischen Reste aus dem Karbon von Karl-Marx-Stadt und Hainichen i. S. – Geologie, 1: 673–694. POSCHMANN, M. & SCHINDLER, T. (1997): Ein Neufund von Fayolia sp. (Chondrichthyes: Hai-Eikapsel) mit paläoökologischen Anmerkungen zum Fundhorizont (Niedermoschel-Bank, Unteres Rotliegend; Saar-Nahe- Becken). – Mainzer geowissenschaftliche Mitteilungen, 26: 25–36. PRANTL, F. (1933): O rodu Palaeoxyris v Čechách. – Rozpravy II. Tridy České Akademie Věd a Umeni, 43 (12): 1–4, Pl. I. PRESL, D.K. (1838): Restiacites. – In: STERNBERG, K.M. (ed.): Versuch einer geognostisch-botanischen Darstellung der Flora der Vorwelt. Vol. II, part VII–VIII, Prag: 189, Pl. LIX. PRESTWICH, J. (1840): On the Geology of Coalbrook Dale. – Transactions of the Geological Society of London 2nd Series, Vol. V (Part III): 413–495, Pl. XXXVIII and Explanations. PRUVOST, P. (1919): Introduction à l'étude du terrain houiller du Nord et du Pas-de-Calais. La faune continetale du terrain houiller du Nord de la France. – Mémoires pour servir à l'explication de la Carte Géologique détaillée de la France. Imprimerie Nationale, Paris: 439–453, Pl. XXVI. PRUVOST, P. (1922): Description d'un fossile nouveau (Scapellites Cottoni nov. gen. et sp.) du Westphalien supérieur du Couchant de Mons. – Annales de la Société Scientifique de Bruxelles, 42: 150–155. PRUVOST, P. (1930): La faune continentale du terrain houiller de la Belgique. – Mémoires du Musée Royal d'Histoire Naturelle de Belgique, 44: 133–141, Pl. II. QUENSTEDT, A. (1867): Handbuch der Petrefaktenkunde. – Zweite umgearbeitete und vermehrte Auflage. H. Laupp'sche Buchhandlung, Tübingen: 982. RANTZIEN, H.H.A. (1956): An annotated check-list of genera of fossil Charophyta. – Micropaleontology, 2 (3): 243–256. RENAULT, B. & ZEILLER, R. (1884): Sur un nouveau genre de fossiles végétaux. – Comptes rendus hebdomadaires des séances de l'Académie des sciences: 1391–1394.

88 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

RENAULT, B. & ZEILLER, R. (1888): Sur l'attribution des genres Fayolia et Palaeoxyris. – Comptes rendus hebdomadaires des séances de l' Académie des sciences, 107: 1022–1025, Pl. XLI–XLII. ROMANOWSKI, G. (1880): Materialien zur Geologie von Turkestan. – I. Lieferung. Buchdruckerei der Kaiserlichen Akademie der Wissenschaften, St. Petersburg: 135–136, Pl. XXIII. RONCHI, A. & SANTI, G. (2003): Non-marine biota from the Lower Permian of the central Southern Alps (Orobic and Collio basins, N Italy): a key to the paleoenvironment. – Geobios, 36: 749–760. RÖSSLER, R. & SCHNEIDER, J.W. (1997): Eine bemerkenswerte Paläobiocoenose im Unterkarbon Mitteleuropas - Fossilführung und Paläoenvironment der Hainichen-Subgruppe (Erzgebirge-Becken). – Veröffentlichungen Museum für Naturkunde Chemnitz, 20: 5–44. SALFELD, H. (1907): Fossile Land-Pflanzen der Rät- und Juraformation Südwestdeutschlands. – Palaeontographica, 54: 164–203. SAMPELAYO, P.H. (1946): Estudios acerca del Carbonifero en España. – Boletín del Instituto Geológico y Minero de España, 59: 1–19, Pl. 2. SANTI, G. (2007): Variation in the Ichnofauna of the Collio Formation (Lower Permian) in the South-Alpine Region (Northern Italy). – Ichnos, 14: 91–104. SAPORTA, G. (1891): Plantes jurassiques. – Paléontologie française ou description des fossiles de la France. Série 2 - Végétaux T 4. Paris: 28–62 & 274–280, Pl. IV–VII. SAUVAGE, H.E. (1905): Note sur un Spirangium du calcaire lithographique de la province de Lérida (Catalogne). – Annales de la Société Géologique du Nord, 34: 9–12. SCHENK, A. (1864): Beiträge zur Flora des Keupers und der rhaetischen Formation. – Siebenter Bericht der naturforschenden Gesellschaft zu Bamberg: 107–108. SCHENK, A. (1867): Die fossile Flora der Grenzschichten des Keupers und Lias Frankens. – C. W. Kreidel's Verlag, Wiesbaden: 149, 195–198, 204–205. SCHENK, A. (1871): Beiträge zur Flora der Vorwelt. IV. Die Flora der nordwestdeutschen Wealdenformation. – Paleontographica, 19: 246–247, Pl. XL–XLI. SCHENK, A. (1888): Die fossilen Pflanzenreste. – Verlag von Eduard Trewendt, Breslau: 185–188. SCHIMPER, W.P. (1870–1872): Traité de paléontologie végétale ou la flore du monde primitif dans ses rapports avec les formations géologiques et la flore du monde actuel. – 2. Bailliére, Paris: 514–519, Pl. LXXX. SCHIMPER, W.P. & MOUGEOT, A. (1844): Monographie des plantes fossiles du grès bigarré de la chaine des Vosges. – G. Engelmann, Leipzig: 46–47, Pl. XXIII. SCHIMPER, W.P. & SCHENK, A. (1890): Palaeophytologie. – In: ZITTEL, K. A. (ed): Handbuch der Palaeontologie, II. Abtheilung. – R. Oldenburg Verlag, München und Leipzig: 393–394. SCHMIDT, F.A. (1855): Petrefactenbuch. Allgemeine und besondere Versteinerungskunde mit Berücksichtigung der Lagerungsverhältnisse, besonders in Deutschland. – Verlag von Krais & Hoffmann, Stuttgart: 39. SCHMIDT, M. (1928): Die Lebewelt unserer Trias. – Hohenlohe'sche Buchhandlung Ferdinand Rau, Öhringen: 47–48. SCHNEIDER, J.W. (1986): Limnoselache n. g. – ein Süßwasserhai der paläozoisch-mesozoischen Familie Tristychiiade (Hybodontoidea). – Freiberger Forschungshefte, C 410: 77–87. SCHNEIDER, J.W. (1996): Xenacanth teeth - a key for taxonomy and biostratigraphy. – Modern Geology, 20: 321–340. SCHNEIDER, J.W., GAITZSCH, B.G., HOFFMANN, U., JASCHKE, I. & WALTHER, H. (2003): Lacustrine Laichbiotope paläozoischer Haie - Reproduktionsmuster mariner Vorfahren? – 73. Jahrestagung der Paläontologischen Gesellschaft 29.9.–3.10.2003, Johannes Gutenberg-Universität Mainz, Terra Nostra - Schriften der Alfred- Wegener-Stiftung (5): 144–145. SCHNEIDER, J.W., HOTH, K., GAITZSCH, B., BERGER, H.J., STEINBORN, H., WALTER, H. & ZEIDLER, M.K. (2005): Carboniferous stratigraphy and development of the Erzgebirge Basin, East Germany. – Zeitschrift der Deutschen Gesellschaft für Geowissenschaften, 156 (3): 431-466. SCHNEIDER, J.W. & REICHEL, W. (1989): Chondrichthyer-Eikapseln aus dem Rotliegenden (Unterperm) Mitteleuropas - Schlußfolgerungen zur Paläobiologie paläozoischer Süßwasserhaie. – Freiberger Forschungshefte, C 436: 58–69. SCHNEIDER, J.W., SIEGESMUND, S. & GEBHARDT, U. (1984): Paläontologie und Genese limnischer Schill- und Algenkarbonate in der Randfazies der kohleführenden Wettiner Schichten (Oberkarbon, Stefan C) des NE- Saaletroges. – Hallesches Jahrbuch für Geowissenschaften, 9: 35–51. SCHULTZE, H.-P. (1995): Terrestrial biota in coastal marine deposits: fossil-Lagerstätten in the Pennsylvanian of Kansas, USA. – Palaeogeography, Palaeoclimatology, Palaeoecology, 119: 255–273. SCHULTZE, H.-P., MAPLES, C.G. & CUNNINGHAM, C.R. (1993): The Hamilton Konservat-Lagerstätte: Stephanian terrestrial biota in a marginal-marine setting. – Transactions of the Royal Society of Edinburgh: Earth Sciences, 84: 443–451.

89 J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91. ______

SCHULTZE, H.-P. & SOLER-GIJÓN, R. (2003): Gedanken zur Paläoökologie europäischer Permo-Karbon-Becken. – 73. Jahrestagung der Paläontologischen Gesellschaft 29.9.–3.10.2003, Johannes Gutenberg-Universität Mainz, Terra Nostra - Schriften der Alfred-Wegener-Stiftung, 5: 150–151. SCHULTZE, H.-P. & SOLER-GIJÓN, R. (2004): A xenacanth clasper from the ?uppermost Carboniferous - Lower Permian of Buxières-les-Mines (Massif Central, France) and the palaeoecology of the European Permo- Carboniferous basins. – Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen, 232 (2/3): 325– 363. SELDEN, P.A. & NUDDS, J.R. (2007): Chapter 7 - Voltzia Sandstone. – In: SELDEN, P.A. & NUDDS, J.R. (eds.): Evolution of Fossil Ecosystems. Manson Publishing Ltd, London: 71–78. SEWARD, A.C. (1894): A new British Carboniferous Fossil. – The Naturalist: 233–240, Pl. I. SOLER-GIJÓN, R. (1997): New discoveries of xenacanth sharks from the Late Carboniferous of Spain (Puertollano Basin) and Early Permian of Germany (Saar-Nahe Basin): Implications for the phylogeny of xenacanthiform and anacanthous sharks. – Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen, 205 (1): 1–31. SOLER-GIJÓN, R. & POYATO-ARIZA, F.J. (1995): Overview of the Early Cretaceous Chondrichthyan fauna from Montsec (Lérida, Spain). – II International Symposium on Lithographic Limestones. Lleida-Cuenca (Spain, 9th–16th July 1995). Extended Abstracts: 145–149. SROKA, S.D. & RICHARDSON JR., E.S. (1997): Problematica. – In: SHABICA, C.W. & HAY, A.A. (eds.): Richardson's Guide to the Fossil Fauna of Mazon Creek. Northeastern Illinois University, Chicago: 270–274. STAINIER, X. (1894): Un Spiraxis nouveau du Devonien Belge. Bulletin de la Société Belge de Géologie, de Paléontologie et d'Hydrologie VIII: 23–28. ŠTAMBERG, S. & ZAJÍC, J. (2008): Chondrichthyan egg capsules. – In: Carboniferous and Permian faunas and their occurrence in the limnic basins of the Czech Republic. Museum of Eastern Bohemia, Hradec Králové: 139. STERZEL, J.T. (1918): Die organischen Reste des Kulms und Rotliegenden der Gegend von Chemnitz. – Abhandlungen der mathematischen-physikalischen Klasse der königlich sächsischen Gesellschaft der Wissenschaften, XXXV (V): 205–315, Pl. I. STIEHLER, A.W. (1850): Herr Stiehler an Herrn v. Carnall. – Zeitschrift der Deutschen geologischen Gesellschaft, II. Band: 181–184, Pl. VII. STIEHLER, A.W. (1860): Die Bromeliaceen der Vorwelt. – Berichte des Naturwissenschaftlichen Vereines des Harzes zu Blankenburg: 4-9. SUTCLIFFE, W.H. (1909): Notes on Palaeoxyris prendeli (LESQUEREUX) from the Middle Coal Measures, Sparth, Rochdale. – Lancashire Naturalist: 117, Text-Fig. 3 on p. 116. SZE, H.C. (1954): Description and discussion of a problematic organism from Lingwu, Kansu, Northwestern China. – Acta Palaeontologica Sinica, 2 (3): 315–322, Pl. I. UNGER, F. (1845): Synopsis Plantarum Fossilium. Lipisiae, Leipzig: 168 V. HAUER, R. (1851): Verzeichniss der an die k. k. geologische Reichsanstalt gelangten Einsendungen von Mineralien, Petrefacten, Gebirgsarten u.s.w. – Jahrbuch der kaiserlich-königlichen Geologischen Reichsanstalt, II. Jahrgang: 156–157. VAN DER HEIDE, S. (1943): La Faune Ichthyologique du Carbonifère Supérieur des Pays-Bas. – Mededeelingen van de Geologische Stichting C-IV-3 (No. 2): 4–65, Pl. 4. VIALOV, O.S. (1984): Nowaja nachodka jaizewych kapsul elasmobranchij Palaeoxyris [New find of elasmobranch egg capsules Palaeoxyris]. – In: VIALOV, O.S., GAWRILISCHIN, W.I., DIDIKOVSKI, W.J., SOSIMOVITSCH, W.J., MAKRIDIN, W.P., MASLUN, N.W. & EYNOR, O.L. (eds.): Paleontologija i stratigrafija fanerosoja Ukrainy [Palaeontology and stratigraphy of the Phanerozoic of Ukraine]. Naukova dumka, Kiev: 99–103 [in Russian]. VOIGT, S., SPINDLER, F., FISCHER, J., KOGAN, I. & BUCHWITZ, M. (2007): An extraordinary lake basin - the Madygen fossil lagerstaette (Middle to Upper Triassic, Kyrgyzstan, Central Asia). – 77. Jahrestagung der Paläontologischen Gesellschaft, Freiberg, 17.–19. September 2007, abstracts; Wissenschaftliche Mitteilungen des Institutes für Geologie, 36: 162–163. WATERLOT, G. (1934): II. Faune Fossile - étude de la faune continentale du terrain Houiller Sarro-Lorrain. – Études des Gîtes Minéraux de la France. Bassin Houiller de la Sarre et de la Lorraine. Lille: 240–244, Pl. XXII. WEISS, C.E. (1884): Beiträge zur fossilen Flora. III. Steinkohlen-Calamarien. II. Nachtrag zu S. 152. Gyrocalamus = Fayolia. – Abhandlungen zur geologischen Specialkarte von Preussen und den thüringischen Staaten, 5 (2): 202–204, Pl. IV. WEISS, C.E. (1887): Ueber Fayolia sterzeliana n. sp. – Jahrbuch der königlich preussischen geologischen Landesanstalt: 94–99, Pl. IV. WHITE, D. (1899): Fossil Flora of the Lower Coal Measures of Missouri. – United States Geological Survey, Washington: 274–275.

90 Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula. ______

WILLIAMS, H.S. (1887): On the fossil faunas of the Upper Devonian. The Genesee Section, New York. – Bulletin of the United States Geological Survey, 41: 86-87. ZAJÍC, J. (1988a): Recent results of the study of Permo-Carboniferous Vertebrates from boreholes in Bohemian limnic basins. – Symosium on Results of the Research on the Permocarboniferous Fauna, Hradec Králové: 11–12. ZAJÍC, J. (1988b): Vertebrates from boreholes in Permo-Carboniferous limnic basins of the Bohemian Massif. – Rotliegendes Lacustrine Basins Workshop, Ksiaz Castle: 1–2. ZAJÍC, J. (2006): The main fish communities of the limnic Permian and Carboniferous basins of the Czech Republic. – 7th Paleontological Conference, Extended abstracts, Scripta Facultatis Scientarum Naturalium Universitatis Masarykianae Brunensis. 33–34 (2003–2004): 99–101. ZAJÍC, J. (2007): Upper Carboniferous non-marine euselachiids of the Czech Republic. – Ichthyolith Issues Special Publication, 40th Anniversary Symposium on Early Vertebrates/Lower Vertebrates, Abstracts, 10: 97–98. ZAJÍC, J. (2008): The main Late Carboniferous and Early Permian lake fish communities of the Czech Republic. – 5th Symposium on Permo-Carboniferous Faunas, July 7.–11. 2008, Museum of Eastern Bohemia at Hradec Králové, Special Publication "Faunas and palaeoenvironments of the Late Palaeozoic": 53–54. ZEILLER, R. (1890): Note rectificative sur le genre Fayolia. – In: RENAULT, B. & ZEILLER, R. (eds.): Etudes sur le terrain houiller de Commentry. Flore fossile. Appendice à la première partie. 4, III série, Bulletin de la Société de l'Industrie Minérale, Saint-Etienne: 369–379. ZESSIN, W. (2008): Einige neue Insekten aus der Unteren Trias (Buntsandstein) von Mallorca, Spanien (Blattaria, Coleoptera, Diptera, Heteroptera und Megaloptera). – Deutsche Gesellschaft für allgemeine und angewandte Entomologie – Nachrichten, 22 (1): 20–21. ZIDEK, J. (1976): A new shark egg capsule from the Pennsylvanian of Oklahoma, and remarks on the chondrichthyan egg capsules in general. – Journal of Paleontology, 50 (5): 907–915.

Paläontologie, Stratigraphie, Fazies Freiberger Forschungshefte, Reihe C

Manuskripte an / send manuscripts to: Prof. J.W. Schneider / Dr. O. Elicki TU Bergakademie Freiberg, Geologisches Institut, D-09596 Freiberg [email protected] / [email protected]