Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula as subject of palaeontological research – an annotated bibliography

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.

Full text

75
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; j.fischer1@yahoo.de
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

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parataxonomische Systematik der Kapseln bleibt problematisch. Die bisher vollständigste Bibliographie zu
diesem Thema schließt diesen Beitrag ab.
1 Introduction
The enigmatic fossil capsules Palaeoxyris B
RONGNIART, 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.

Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula.
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77
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

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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.

Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula.
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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.

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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 &

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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.

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82
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.

Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula.
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83
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 &

J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91.
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84
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).

Elasmobranch egg capsules Palaeoxyris, Fayolia and Vetacapsula.
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85
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 intra-
specific 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.

J. Fischer & I. Kogan / Freiberger Forschungshefte C 528 – psf 16 (2008): 75 – 91.
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Paläontologie, Stratigraphie, Fazies
Freiberger Forschungshefte, Reihe C
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Manuskripte an / send manuscripts to: Prof. J.W. Schneider / Dr. O. Elicki
TU Bergakademie Freiberg, Geologisches Institut, D-09596 Freiberg
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