PalZ (2019) 93:207–253

https://doi.org/10.1007/s12542-018-0433-5

RESEARCH PAPER

Helcionelloids, stenothecoids and hyoliths from the Tannenknock

Formation (traditional lower middle Stage 4/Wuliuan boundary
interval) of the Franconian Forest, Germany
Gerd Geyer1   · Martin Valent2 · Stefan Meier3

Received: 23 December 2017 / Accepted: 8 August 2018 / Published online: 9 November 2018
© Paläontologische Gesellschaft 2018

Abstract
The early middle Cambrian Tannenknock Formation of the Franconian Forest, traditionally subdivided into the Galgen‑
berg and Wildenstein strata, yields a relatively diverse fauna. This fauna includes a number of molluscs, which portrays a
fairly diverse assemblage that can be regarded as paradigmatic for the lower–middle Cambrian boundary interval in West
Gondwana, with the helcionelloid genera Helcionella, “Bemella,” “Igorella,” Latouchella, Leptostega, Yochelcionella, the
pelagiellid Pelagiella, a stenothecoid, and the Hyolithida Hyolithes?, Hexitheca?, Jincelites, Maxilites?, Aladraco, Cam-
brachelous, the Orthothecida Brevitheca?, Nephrotheca?, and a number of helcionelloid and hyolith forms dealt with under
open nomenclature. New species include the helcionelloids Helcionella lemdadensis sp. nov. and Leptostega frankenwalden-
sis sp. nov. and the hyoliths Grantitheca? klani sp. nov. and Cambrachelous diploprosopus gen. et sp. nov. The stenothecoid
discovered in the Wildenstein Member of the formation is the first report of the group from West Gondwana.

Keywords  Cambrian · Helcionelloida · Stenothecoida · Hyolithida · Biostratigraphy · West Gondwana · Germany

Introduction Wurm’s initial publications (Wurm 1924a, b, 1925a, b) pro‑

vided preliminary data on the fossils. Wurm recognised an early
The Cambrian of the Franconian Forest area (German: Frank‑ Middle Cambrian fauna with trilobites, brachiopods, hyoliths
enwald) in northeastern Bavaria is among the oldest known and echinoderm remains, which were subsequently attributed
areas with Cambrian rocks in Europe outside Scandinavia and to two different formations, called Galgenberg and Wildenstein
the British Isles, but remained largely unstudied after its first strata, respectively (Sdzuy 1964). Wurm (1925b) recognised
discovery during a mapping season in 1923 by Adolf Wurm. three hyoliths species, which were dealt with in open nomencla‑
ture as Hyolithes sp. a, Hyolithes sp. b, and Hyolithes sp. c. In
addition, he described three species attributed to Conularia as
Handling editor: Mike Reich.
Conularia sp. a, C. sp. b, and C. sp. c and introduced a new spe‑
* Gerd Geyer cies under the name “Conularia Schloppensis”, later transferred
gerd.geyer@uni‑wuerzburg.de to the new genus Oxyprymna Kiderlen, 1933 (now Aladraco
Martin Valent Geyer, 2018). In fact, all of them represent hyoliths. Helcionel‑
martin_valent@nm.cz loid molluscs were not reported to date, but are known to be
Stefan Meier represented in a large number in rocks of the Tannenknock For‑
stefan.meier.mak@t‑online.de mation and particularly in the Wildenstein Member and are
1
described below for the first time.
Institut für Geographie und Geologie, Lehrstuhl für
Geodynamik und Geomaterialforschung, Bayerische
Julius-Maximilians-Universität Würzburg, Am Hubland,
97074 Würzburg, Germany Geological setting and stratigraphy
2
Národní Muzeum, Václavské náměstí 68, 115 79 Praha 1,
Czech Republic The Franconian Forest lies on the margin of the Saxo‑
3
Mineralienkabinett Stefan Meier, Zweigstraße 22, thuringian Zone in northeastern Bavaria, southern
95615 Marktredwitz, Germany Germany (Fig.  1). The Saxothuringian Zone is a West

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208
Fig. 1  Generalized geological map of Saxothuringian Zone in north‑
eastern Bavaria, southern Germany; small-scale insert map shows
modern political boundaries of Germany and adjacent regions with
outcrop areas of Cambrian rocks (black), outcrop areas of rocks
Gondwana-associated terrane or, better, a marginal West
Gondwana succession (Geyer et al., in rev.). The Cam‑
brian–Carboniferous successions of the Saxothuringian
Zone are now interpreted as giant olistoliths in wildfly‑
sch of the Variscan orogen (e.g. Linnemann and Schauer
1999). Cambrian rocks in the Saxothuringian Zone show
a geographically coherent facies succession that does not
suggest either regional allochthony or transport for most of
the zone (e.g. Göthel 2001). However, the small Cambrian
blocks in the Franconian Forest are obviously related to the
presence of the so-called Münchberg Gneiss Mass, a genet‑
ically and chronologically controversely debated block of
metamorphic rocks (e.g. Stettner 1972; Behr et al. 1980;
Gandl 1998) (Figs. 1, 2).
13
G. Geyer et al.
with supposed Cambrian portions (grey), and subsurface Cambrian
(hatched). Small white rectangle outlines refer to the tectonic units
with Cambrian strata: A, Wildenstein slice (detailed map in Fig. 2); B,
Triebenreuth slice
The Franconian Forest has a middle Cambrian succes‑
sion of siliciclastic-dominated, shallow-water, fossiliferous
units. The formations of early middle Cambrian age are of
typical West Gondwanan aspect, similar to that described
from the Moroccan Atlas ranges (Geyer and Landing 1995,
2006; Geyer et al. 1995). This succession has primarily West
Gondwanan trilobites in its lower part (traditionally termed
Galgenberg and Wildenstein “strata”, or formations) (e.g.,
Geyer et al. 2008; Heuse et al. 2010; Geyer 2017).
Wurm’s early publications on these strata (Wurm 1924a,
b, 1925a, b) recognised an early Middle Cambrian (in tra‑
ditional sense) fauna from probably four adjacent localities
in the vicinity of the so-called Galgenberg (“Gallow Hill”)
near Wildenstein. This fauna turned out to include fossils
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
Fig. 2  Areas with outcrops of Cambrian rocks of the Wildenstein
slice. Based on maps of Wurm (1925b), Sdzuy (1964), Ludwig
(1969), Horstig and Stettner (1976), and new data. Brown colour
marks outcrop areas dominated by Galgenberg facies, blue colour
outcrop areas dominated by rocks of the Wildenstein Member. Num‑
bers refer to exposures or sampling sites (see “Appendix”)
attributable to two differentiable stratigraphic horizons. He
referred to the units in the sense of formations, which were
formally named by Geyer and Wiefel (1997). Typical fossils
from the Galgenberg facies occur in light coloured sandy,
209
slightly calcareous blue-grey shales, which tend to weather
with whitish outer surfaces, whereas Wildenstein fossils can
be found in fine- to medium-grained, slightly calcareous,
commonly feldspathic sandstones or ochre-weathering sandy
calcareous nodules (Geyer et al. 2008; Geyer 2010, 2017).
Recent investigations suggest that both units represent in
fact a single formation with temporal facies shifts that cul‑
minate in the typical Galgenberg and Wildenstein facies.
This formation is formally introduced and dealt with as the
Tannenknock Formation (Geyer 2017). The Tannenknock
Formation with both facies expressions (now separated as
members) was traditionally known from just two isolated
blocks, termed the Wildenstein (prefix “W” in the locality
IDs) and Triebenreuth (prefix “T” in the locality IDs) slices.
The vast majority of the material studied herein comes from
the Wildenstein slice. The geological situation of this Wil‑
denstein slice has been studied in detail by Ludwig (1969)
and remapped in the course of this study (Fig. 2).
Age and correlation
Despite the limited amount of source rocks, the trilobite
faunas from the Galgenberg and Wildenstein members of
the Tannenknock Formation are comparatively diverse and
enable a precise correlation with other Cambrian areas of
West Gondwana (Fig. 3). Particularly similar are the trilobite
faunas from the Jbel Wawrmast Formation of the Moroccan
Atlas ranges. Characteristic for the Galgenberg Member is
Kingaspidoides frankenwaldensis (Wurm, 1925a, b), a spe‑
cies that has been found in the Moroccocus notabilis Zone
(earlier the Cephalopyge notabilis Zone) of the eastern Anti-
Atlas (Geyer and Vincent 2015). Other faunal elements of
the Galgenberg Member have closely related counterparts in
the same zone in the Atlas ranges. The trilobite associations
of the Wildenstein Member is characterised by Ornamen-
taspis frequens Geyer, 1990, a species that is an index fos‑
sil for the O. frequens Zone in the Moroccan Atlas ranges,
which directly overlies the M. notabilis Zone, thus providing
a perfect match for the formations in the Franconian Forest.
Rocks from near Wustuben (ca. 4.3 km NNE of the northern
tip of the Wildenstein slice (ca. N 50° 15′ 0″, E 11° 34′ 20″)
newly discovered by S. Meier also include numerous hel‑
cionelloids. They belong to a succession superficially similar
to those of the Wildenstein Member of the Tannenknock
Formation, but with a considerable content of volcaniclastic
material. According to the trilobite fauna, they come from
a hitherto unknown younger stratum of late Agdzian–early
Caesaraugustan assigned to the overlying Triebenreuth
Formation (Fig. 3). The helcionelloids from these strata are
similar to those of the Tannenknock Formation and will be
descrbed separately except for “Igorella” sp. A (see below).
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210
Fig. 3  Stratigraphic table of the fossiliferous strata in the Franconian
Forest with highlighted Tannenknock Formation and its correlation
with global chronostratigraphic units and the standard Cambrian of
West Gondwana (Geyer and Landing 2004)
Of the helcionelloids described herein, Helcionella
capula Geyer, 1986 has been described from the Moroc-
conus notabilis Zone of the Jbel Wawrmast Formation in
the Lemdad area, High Atlas, and specimens tentatively
assigned to the species occur in the O. frequens Zone of the
Lemdad Syncline. Leptostega was originally described from
lower middle Cambrian strata of the Cantabrian Mountains,
Spain (Geyer 1986), but was subsequently recognized from
a number of other regions. However, the generic concept of
Geyer (1986) has been misinterpreted (see below under Lep-
tostega) so that the genus is momentarily known from West
Gondwana, eastern Avalonia and the Siberian Platform only.
Remarkably, similarities can be recognized with helcionel‑
loid faunas known from the Doberlug region of northern
Saxony (Schmidt 1942), but less so with such known from
the Görlitz region of Lusatia (e.g., Elicki 2003, 2007).
It is remarkable that the helcionelloid fauna now recov‑
ered from the overlying late Agdzian–early Caesaraugus‑
tan strata of the Triebenreuth Formation (to be described
subsequently) is fairly similar to that of the early Agdzian
Tannenknock Formation and even has species/forms in com‑
mon. This suggests that the occurrence of Helcionelloida is
largely determined by facies and of little biostratigraphic
significance, at least in this chronostratigraphic interval.
Aladraco schloppensis, found in the Franconian Forest in
the Galgenberg Member and dominantly in the Wildenstein
Member of the Tannenknock Formation, has a counterpart
with Aladraco ougnatensis from the upper part of the M.
notabilis Zone in the eastern Anti-Atlas (Geyer 2018). The
hyolith fauna described herein is remarkably diverse and
13
G. Geyer et al.
includes forms assigned or tentatively assigned to the hyo‑
lithid genera Hyolithes, Hexitheca, Jincelites, Maxilites,
and Aladraco and Cambrachelous, the orthothecid genera
Brevitheca, Nephrotheca, as well as forms that cannot be
assigned to a genus with any confidence. The reason for the
problems of a definite assignment is caused by an imperfect
preservation or the absence of both conchs and opercula,
but also by the incomplete knowledge on hyoliths from
this stratigraphic interval of the late part of the Cambrian
Stage 4 and the early part of Stage 5 in West Gondwana and
elsewhere.
Localities and collections
Because of the extremely small outcrop areas and dense for‑
ested vegetation or agricultural use of these areas, collect‑
ing Cambrian fossils is strenuous or depends on fortuitous
conditions. The fossils used in this and other publications are
exclusively part of the fauna of the Galgenberg and Wilden‑
stein members of the Tannenknock Formation, originating
from the Wildenstein slice (Fig. 1a) and the Triebenreuth
slice (Fig. 1b). The exact locations of the sampling sites are
shown in Fig. 2 with additional data provided in “Appendix”.
Further data in the lists of material in the systematic section
refer to those localities.
Collecting started with A. Wurm’s discoveries in the
1920s. A wealth of samples originate from unstudied col‑
lections of K. Sdzuy (mostly from 1956), collections of
unknown origin left in the former Institut für Paläontologie,
Würzburg University (abandoned 2004/2005), and collec‑
tions of the senior author (1979‒2013). An unexpectedly
large amount of additional specimens comes from recent
collections by Stefan Meier (see “Appendix” for further
information).
Regrettably, the knowledge of the exact location of col‑
lecting sites most of the early collections during the early
stage of research in the region, particularly for those of
Adolf Wurm, S.G. Kohlmann and Hermann Klan is sparse
to non-existent. The material collected by K. Sdzuy comes
from a number of different localities in the Wildenstein slice,
which fortunately could be confidently located based on his
field notes of 1956.
Preservation
The preservation of the molluscs described herein depends
on the facies and preservational condition within the host
rock. Specimens from the Wildenstein Member are usu‑
ally found in calcareous arenites or large calcareous nod‑
ular-type concretions (of up to meter-size), from which the
carbonate has been largely dissolved by weathering so that
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
the specimens are found as internal moulds and/or casts of
the exterior. This provides three-dimensionally preserved
specimens often with minor distortion. However, fine details
of the surface ornament are rarely preserved under these
circumstances. The fauna includes most of the helcionel‑
loid specimens and the pelagiellids, but hyoliths are usu‑
ally absent from these strata. Shaly rocks of the Wildenstein
Member may include large specimens of hyoliths, such as
Aladraco schloppensis. The specimens from these strata are
moderately to strongly flattened and preserved in different
ways, including composite moulds.
The sandy shales of the Galgenberg Member yield rela‑
tively few hyoliths, in which the shell is preserved in various
modes of pseudomorphy. Specimens in which the original
shell material was replaced by ferritic minerals, now vis‑
ible as a yellowish or reddish substance, are common and
this also includes the remineralisation of apical septa. Some
portions include hyolith specimens (usually orthothecids) in
which the shell material is now phosphatised. These fairly
common specimens are often recognisable in the rock by
their elliptical cross-sections. Phosphatisation unfortunately
did not just replace the shell, but the mineral growth pro‑
cess continued so that fine details of the exterior are rarely
preserved.
A particularly noteworthy fact is that a number of hyo‑
lith conchs exhibit a selective phosphatisation in which the
phosphatic steinkern can be observed within the hyolith shell
(Figs. 13a, f, l, 14g, 15e, 23b). As seen in other Cambrian
shelly fossils, small cavities appear to provide geochemical
conditions that favoured a reprecipitation of apatite, which in
turn made possible the extraction of millimetre-sized “Small
Shelly Fossils” by dissolving early Cambrian carbonates,
resulting in a strong taphonomic bias toward small-sized
fossils (Creveling et al. 2014). Phosphatisation in these cases
took place early in the diagenetic process because details of
the internal shell surface are preserved with fidelity prior to
the dissolution of the shell (e.g., Vendrasco et al. 2010). In
the case of the Wildenstein hyolith, the phosphatic partial
steinkerns are incomplete in a way that reflects the limits of
the geochemical conditions ruling within the apical parts of
the hyolith conchs.
Ecological aspects
Helcionelloida and Stenothecoida
The synecology of the Helcionelloida in the Tannenknock
Formation primarily reflects environmental constraints.
Fairly frequent occurrences of these fossils in the large cal‑
careous sandy nodules of the Wildenstein Member (such as
in sample W8) resulted from favourable conditions both in
biotic association and taphonomic history. The abundance
211
of helcionelloid specimens is unambiguously controlled by
winnowing and size sorting of small sclerites, or fragments
of sclerites, during the sedimentation of the wave-controlled
shell accumulations, which form the major part of the fossil‑
iferous calcareous strata of the Wildenstein Member facies.
The particular composition of the richly diverse associations
differs moderately between the sample horizons, which is
remarkable because of the short distance between the locali‑
ties. It may be assumed that the composition of the same
stratum may have changed dramatically over considerable
distance, similar to what can be observed for the Brèche
à Micmacca limestone beds in the Moroccan Atlas ranges
(Geyer and Landing 1995; Geyer et al. 1995).
Because of the wave action during deposition, larger
shells and sclerites are usually crushed, but preservation of
sharp edges and absence of abrasion excludes the possibil‑
ity of notable transport. In total, only about 90 specimens
of helcionelloids have been found in the studied samples of
which ca. one third is preserved as small fragments only.
Larger helcionelloid shells are often preserved more or less
intact, but this is certainly a result of favourable transport
properties of the small conical conchs.
The number of specimens in most samples is regarded
as insufficient to allow calculation of robust statistical
data. Eight helcionelloid species and recognizable forms,
one pelagiellid and one stenothecoid are described below.
Nevertheless, for the samples with the highest number of
helcionelloid specimens (sample horizon W8), the helcionel‑
loids contribute to less than 0.5% of the total fauna in terms
of specimens. The total fauna from the same strata registered
to date includes at least 14 different trilobites, five brachio‑
pods, at least five different echinoderms, plus several other
faunal elements (such as sponges, palaeoscolecidans, and
various echinoderm groups). This reflects a composition
similar to what is known from more or less coeval strata
in other Cambrian regions (such as the Moroccan Atlas
regions; Geyer and Landing 1995; Geyer et al. 1995).
Hyolitha
Hyoliths are favourably preserved in shaly strata of the Wil‑
denstein facies which indicates relatively quiet depositional
conditions. As a result, the hyoliths conchs are completely
embedded usually although notable bioturbation may be
observed in those rocks. Nevertheless, rare cases of speci‑
mens with tips of the conchs broken off are now recorded,
and some specimens show broken apertural margins.
Taphonomic and diagenetic effects led to preservation of
specimens as composite moulds with a notable dorsoven‑
tral compaction of the conchs. Those specimens may show
fractures along the sagittal/exsagittal axes of the distinctly
convex parts of the conchs, but this is not always the case.
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212
The shaly rocks of the Wildenstein facies includes a com‑
paratively paucispecific faunal assemblage with only ca. four
trilobite species (including Acadoparadoxides, Kingaspi-
doides, and Bailiella) and the eight hyolith species/forms
described below as regular faunal elements.
Systematic palaeontology
The type material and the figured specimens used in this
study are reposited in the collections of the Bayerisches
Landesamt für Umwelt (LfU), the Bayerische Staatssa‑
mmlung für Paläontologie und Geologie, München (SNSB-
BSPG and PIW), and the collection Stefan Meier, Mark‑
tredwitz (SSMM), respectively, under the listed collection
number.
For most of the sampling localities, it appears to be dif‑
ficult to collect additional specimens under standard con‑
ditions so that no further description of material can be
expected during next few decades. Therefore, even imper‑
fectly preserved specimens are described in some detail in
the systematic section because they provide important infor‑
mation on the faunal spectrum.
Phylum Mollusca Cuvier, 1797
Class Helcionelloidea Peel, 1991
Emended diagnosis. Generally bilaterally symmetrical uni‑
valves in which the calcareous shell is usually coiled through
up to several whorls; the whorls may be in contact or open
coiled and are often laterally compressed. The aperture is
oval, without re-entrant, but the sub-apical surface may
develop a median sinus which in some taxa is deep and slit-
like or trematose, with a single perforation at the end of an
elongate tube termed the snorkel. In some forms, the lateral
areas of the aperture may become prosocyrt, extended into
weak lateral fields and producing broad emarginations in
both the supra-apical and sub-apical surfaces astride the
plane of symmetry. Ornamentation may include both comar‑
ginal and spiral elements; prominent comarginal rugae are
common (modified from Peel 1991).
Remarks. Helcionelloid molluscs are among the most
frequent and well-studied components of the Cambrian
“Small Shelly Fossils.” However, the number of publica‑
tions strongly contrasts with evidence for their anatomical
organization and phylogenetic pathways. Consequently, a
number of different approaches for taxonomic subdivisions
have been applied, particularly at family level. Use of the
taxon “Monoplacophora” by various workers following
the introduction of the classes Helcionelloidea (Peel 1991)
and Tergomya (Horný 1965) reflects a lack of insight into
the taxonomy of this group. Parkhaev (2001, 2008) and
13
G. Geyer et al.
subsequent publications, (e.g., Parkhaev 2017a) interpreted
the helcionelloids as belonging to the Class Gastropoda and
representing the most ancient already torted univalved mol‑
luscs, initially symmetrical externally, and later gradually
developing its external asymmetry. This is an opinion for
which conclusive data are not apparent and which is not
shared herein.
Jacquet and Brock (2016) investigated a potential cor‑
relation between ontogenetic developments and taxonomic
conditions. Based on silicified macrohelcionelloids from
the lower Cambrian of South Australia, they analysed the
change in growth from the apical part to large, adult shells,
which obviously indicates a transformation from larval
shells via juvenile conchs (if developed) to teleoconchs. This
topic has also been addressed in some detail by Martí Mus
et al. (2008). The changes during ontogeny and their precise
modality provides clues for a more character-based taxo‑
nomic treatment and sheds light on a number of microscopic
forms which represent just juvenile specimens of unknown
macrohelcionelloids. It should be emphasized, however,
that the number of Cambrian micromolluscs exceeds the
number of centimeter-sized specimens by far (probably up
to one hundred times), but there is no common agreement
on whether this is an evolutionary phenomenon or a tapho‑
nomic artefact. In any case, it is certainly true that some of
these abundant micromolluscs represents juveniles, but not
the majority.
It is obvious that the specific morphological characters of
immature individuals reflect living and habitat conditions.
Consequently, feeding strategies should be recognizable by
the morphology of larval shells (e.g., Runnegar 2007). How‑
ever, much more macroscopic material is needed to over‑
come the momentary artificial subdivision based mostly on
small specimens recovered from dissolved samples so that
studies based on insufficient material may prompt remarks
about the “epitome of hand waving” (Rouse 2000).
Order Helcionellida Geyer, 1994
Family Helcionellidae Wenz, 1938
Genus Helcionella Grabau and Shimer, 1909
Type species. Metoptoma? rugosa Hall, 1847 [= Helcion sub-
rugosa (d’Orbigny, 1850)], from the upper part of the lower
Cambrian Browns Pond Formation (formarly Schodack For‑
mation) of the Taconic Allochthon, Troy, New York State.
Emended diagnosis. Small to large, bilaterally symmetri‑
cal univalve Helcionellidae with nearly patelliform to low
cap-shaped, cyrtoconic shell ornamented by wide, strongly
convex concentric plicae and closely spaced ribs, some‑
times radial ribs of about the same size as the concentric
elements. Apex inclined, not projecting to or beyond the
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
apertural margin. Supra-apical surface of the conch more
or less evenly convex, sub-apical surface concave, without
distinct bands. Aperture planar in lateral view, oval or sub‑
circular in plan view; apertural margin without flarings or
notches. (Modified from Jacquet and Brock 2016).
Discussion. The genus Helcionella is generally regarded as
representing the basic plan of a helcionelloid conch, with
more or less simple concentric rugae or plicae, a simple
coiling up to less than half a whorl, and without any modifi‑
cations of the aperture and the sub-apical surface. This form
may occur in miniaturised shape as a juvenile part in shells
of other helcionelloid genera. However, Helcionella has a
comparatively late occurrence among the lower Cambrian
helcionelloids.
First used as a basic collective genus and subsequently as
a waste basket for simple and often imperfectly preserved
helcionelloid conchs, the genus (or at least its primary
morphological concept) has been split during the past few
decades into numerous genera, which are mostly well dis‑
tinguished. A few of these genera, however, are difficult to
precisely separate. Their species may accumulate in clusters
of typical morphologies, but the morphospace between them
is bridged by a number of species which cannot be placed
unequivocally. In addition, so-called diagnostic characters
provided by most authors are often of little biological mean‑
ing (in terms of phylogenetic connotation) and barely of any
significance with respect to soft-part organization. The com‑
mon approach to overcome this deficiencies is to base the
generic concepts on the type species’ morphologies. This,
however, creates random morphospaces and depends on the
haphazard history of fossil discovery rather than a serious
attempt to understandevolutionary developments.
Such genera include Bemella Missarzhevsky in Rozanov
et al. (1969), and Ilsanella Missarzhevsky, 1981. Ilsanella
has been intruduced to replace the Ginella Missarzhevsky in
Rozanov and Missarzhevsky (1966) as a younger homonym
of the Middle Ordovician ostracod genus Ginella (Ivanova
1955). Ilsanella was interpreted to be distinguished from
Helcionella by its usually much higher shell by Parkhaev
(2001) although this author discussed the difficulties to dis‑
cern the genera if intermediate forms (such as Helcionella
capula Geyer, 1986) come into discussion (Parkhaev 2001,
p. 149). The arbitrary distinction between the genera is rec‑
ognizable at species with intermediate heights that do not
provide a diagnostic difference. By contrast, the mode of
coiling appears to be a simple effect of allometric growth
and can be altered easily by a simple variation of any of
the three major distinguishing parameters (e.g., Thompson
1945; Raup 1966; Gould 1968) although this does not attest
that such changes in the shape do not depict valuable evolu‑
tionary processes. However, differentiation based on robust
213
morphologic features does not generally exist at our present
state of knowledge.
Jacquet and Brock (2016) revised the group and sug‑
gested a distinction between Helcionella and Ilsanella
based on a combination of characters. They proposed that
Helcionella is “characterised by a low to moderate profile
… with rapid lateral expansion of the shell resulting in the
presence of bifurcating rugae” and Ilsanella generally lacks
this bifurcation and “has a comparatively taller shell with
H:L ratios of ≥ 0.75 … and a slower rate of coiling and lat‑
eral expansion” so that “most species of Ilsanella develop
distinctive, uniformly rounded rugae that girdle the entire
shell circumference without requiring excessive bifurca‑
tion on the supra-apical surface” (Jacquet and Brock 2016,
p. 351). Indeed, this distinction leads to a practical option
to distribute the species between Helcionella and Ilsanella
in the revised concepts. However, the new concepts do not
reflect the principal differences between the type species,
and, more important, they suggest different phylogenetic
pathways depicted by the species of both genera. Conse‑
quently, Ilsanella Missarzhevsky, 1981 is considered here
a junior synonym of Helcionella Grabau and Shimer, 1909.
The generic diagnosis of Bemella has been emended by
Parkhaev (2001) to include low cap-shaped, moderately
compressed shells with a posteriorly inclined apex and a
simple, plane aperture. Parkhaev (2001, pp. 152‒154)
regarded a projection of the apex beyond the margin in dor‑
sal view as the most diagnostic character to separate Bemella
from Helcionella although a number of species included
in his list have an apex that does not project as far poste‑
riorly. This character is regarded herein as insufficient to
characterize Bemella, and it also has little biological and
phylogenetic consequence and should, therefore, not serve
as a taxonomic criterion. A more important character is the
apparent difference in morphology between the protoconch
or at least juvenile part of the shell and the teleoconch,
which are clearly separated in the typical species of Bemella,
B. jacutica Missarzhevsky in Rozanov and Missarzhevsky,
1966, which is known from well preserved specimens (e.g.,
Rozanov et al. 1969, pl. IV, fig. 3, Rozanov et al. 2010, pl.
27, fig. 6a). These species are also characterized by a slight
lateral compression. However, it needs to be emphasized
that a number of species and forms attributed to Bemella
and showing close morphological resemblance to B. jacutica
are possibly just juvenile stages of shells that belong to other
genera or are just impossible to attribute to one of the genera
of this group. A compulsory discussion of the numerous spe‑
cies previously attributed to Bemella and their taxonomy is
beyond the scope of this study.
Helcionella capula Geyer, 1986
Figure 4a–i, k–m, o–q, j?, n?
13

214
* v Helcionella capula n. sp.—Geyer 1986, p. 72, pl. 1,
figs. 1–7.
Material. 16 (possibly ca. 25) specimens, mostly internal
moulds and composite moulds. From sample locality W8:
MMUW 2014A-110, MMUW 2014A-117, MMUW 2014A-
118, MMUW 2014A-120, MMUW 2014A-122a, MMUW
2014A-159, MMUW 2014A-160, MMUW 2014A-179.
From sample locality W8: MMUW 2014A-124a, MMUW
2014A-126, SSMM 10406, SSMM 11407, SSMM 10421a,
SSMM 10787c, SSMM 11036b, and SSMM 10797a. From
sample locality W13e: MMUW 2014A-142. Tentatively
assigned to H. capula: From sample locality W8: MMUW
2014A-106, MMUW 2014A-108, MMUW 2014A-109,
MMUW 2014A-112, MMUW 2014A-155a, MMUW
2014A-194.
Localities and stratum. Wildenstein slice, Franconian Forest,
sample localities W8 and W13e. Tannenknock Formation,
Wildenstein Member.
Description. Isostrophic shell forms a cap-shaped, faintly
recurved, compressed cyrtoconic conch with a large suboval
to slightly subrectangular aperture. Maximum length of the
studied specimens ranges from ca. 3 to 11 mm, max. width
2.7 to ca. 9 mm, with an average length/width ratio of ca.
10:8 or 10:7. In the early growth stages, the cross-section is
suboval and ornamented by low transverse costae. Initially,
the costae extend around the entire shell in subequal breadth
but gradually become narrower along the sub-apical area so
that their margins describe more and more obvious acute
angles in lateral view and become more and more promi‑
nent as the aperture is approached. Well preserved inter‑
nal moulds show narrow and fine, pseudoseptate incisions,
which are characteristic for H. capula. This feature results
from incised ledges in respect to the faces of the ridged con‑
centric rugae and is termed “angular grooved” by Jacquet
and Brock (2016).
The shell exterior shows a finely reticulate pattern on the
adult conch. The intersections of the longitudinal and trans‑
verse elements within the reticulate pattern appear to create
small and low nodes. The shell interior shows a similar, but
very fine reticulate pattern that is only visible in well pre‑
served specimens.
Discussion. The present material from the Franconian For‑
est matches the characters seen in the type material from
the High Atlas, Morocco (Geyer 1986). A few specimens
from the Franconian Forest samples extend the sizes seen
in the Moroccan material, and these specimens show more
prominent concentric rugae that are subangular in transverse
section (Fig. 4g, h) on some specimens. The reticulate pat‑
tern in those specimens is that seen in smaller specimens and
13
G. Geyer et al.
frequently includes a subdivision of a concentric rib into a
broader and a much narrower band by a small concentric
lira (Fig. 4i).
Two other forms of Helcionella co-occur with H. capula
in the Wildenstein facies of the Tannenknock Formation.
Helcionella aff. oblonga Cobbold, 1921 is distinguished
by the lack of the typical pseudoseptate incisions and the
possessopm of a distinctly protruding apical portion on
generally larger conchs. Helcionella sp. A also lacks the
pseudoseptate incisions, is generally larger and has a more
subcentrally located apex and somewhat more irregular
plications.
Helcionella lemdadensis sp. nov.
Figure 5n
non * 1921 Heliconella [sic!] oblonga, sp. nov.—Cobbold,
p. 365, pl. XXIV, figs. 38, 39.
v 1986 Helcionella oblonga Cobbold 1921—Geyer,
p. 74–75, pl. 1, fig. 9a–c.
pars 2016 H. oblonga—Jacquet and Brock, p. 350.
Etymology. Named after its occurrence in the Lemdad syn‑
cline, High Atlas, Morocco.
Holotype. SMF 34737, fairly complete internal mould of
conch (Fig. 5n and Geyer, 1986, pl. 1, fig. 9a–c).
Type locality and type stratum. Lemdad Syncline, High
Atlas, Morocco, section Le II, horizon X255; Tatelt Forma‑
tion, Morocconus notabilis Zone, lowermost middle Cam‑
brian (see more information in Geyer et al. 1995).
Diagnosis. Species of Helcionella with patelliform, moder‑
ately high shell, width/length ratio ca. 0.71; apertural outline
subrectangular, with subparallel lateral margins, relatively
straight anterior margin and more regularly curved poste‑
rior margin; apex strongly recurved, blunt, situated ca. 6%
from anterior margin; first order ornament consists of a
maximum of ca. 12 rounded concentric rugae; second order
radial ribs intersect rugae perpendicularly to form distinct
cross-hatched ornament, crossed by much wider standing,
low concentric ribs.
Discussion. Helcionella lemdadensis sp. nov. is based on
only two specimens, one of which was lost during the initial
study by Geyer (1986). Geyer (1986) assigned the specimens
from the High Atlas Mountains, Morocco, to H. oblonga
Cobbold, 1921, the latter species originally described from
the Paradoxides groomi beds (now the lowermost part of the
lower middle Cambrian Upper Comley Sandstone) of Com‑
ley, Shropshire. Helcionella oblonga is comparable in size,
length/width ratio, the type of corrugation and the pattern
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
Fig. 4  a–i, j?, k–m, n?, o–q Helcionella capula Geyer, 1986. a
SSMM 10406, partial conch, internal mould, Tannenknock Forma‑
tion, Wildenstein Member, sample locality W8, oblique view; b, f
SSMM 10421a, partial conch, internal mould, sample locality W8;
dorsal and posterior views; c MMUW 2014A-118, latex cast of
external mould, sample locality W8; posterolateral view; d MMUW
2014A-120, fragment of composite mould, sample locality W8; e
MMUW 2014A-142, fragment of internal mould, sample locality
W13e; g, h MMUW 2014A-159a, latex cast, sample locality W8;
posterior and dorsal views; i MMUW 2014A-122a, fragment of inter‑
nal mould, sample locality W8; ventral view; j, n MMUW 2014A-
106, dorsoventrally compressed conch, internal mould, sample local‑
215
ity W8; dorsal and oblique lateral views; k MMUW 2014A-117,
internal mould, sample locality W8; lateral view; l, m SMF 34729,
holotype, internal mould, Lemdad syncline, High Atlas, Morocco,
section Le XV, horizon 8/2; posterior and lateral views (specimen
figured in Geyer, 1986, pl. 1, fig.  1; o, p MMUW 2014A-160, latex
cast of external mould, slightly dorsoventral compressed, sample
locality St; dorsal and lateral views; q SMF 34876, paratype, Lemdad
syncline, High Atlas, Morocco, section Le I, horizon 2; dorsal view
(specimen figured in Geyer, 1986, pl. 1, fig.  6). All specimens from
Tannenknock Formation, Wildenstein Member, except otherwise
noted. Scale bars 1 mm
13

216
Fig. 5  a–c, d?, e–i, j?, k–m Helcionella aff. oblonga Cobbold, 1921.
a, e MMUW 2014A-148c, incomplete conch, internal mould, sam‑
ple locality W8; lateral and dorsal views; b, f, MMUW 2014A-148a,
incomplete conch, internal mould, sample locality W8; oblique lateral
and dorsal views; c, g, MMUW 2014A-148b, partial conch, inter‑
nal mould, sample locality W8; oblique anterior and dorsal views;
d, MMUW 2014A-152, fragment of conch, external mould showing
reticulate pattern of shell created by delicate costae, sample locality
W8; h, MMUW 2014A-113, external mould, sample locality W8;
i, MMUW 2014A-105, internal mould of incomplete conch, lateral
view, sample locality W8; j, MMUW 2014A-131, fragment of conch,
of reticulate ribs. It has been assumed that the considerably
lower height of the species from Comley (based on a sin‑
gle specimen) can be attributed to dorsoventral compaction.
However, restudy of the holotype of H. oblonga (SM A440
in the Sedgwick Museum, Cambridge), indicates that com‑
paction does not explain the lower height. The species from
Shropshire indeed has a slightly higher width/length ratio of
ca. 0.68 than H. lemdadensis (ca. 0.71) and a slightly more
projecting apex (tip broken off in the holotype).
Another somewhat similar species, Helcionella histosia
Jacquet and Brock, 2016, has been described from the Third
Plain Creek Member of the Mernmerna Formation, Flin‑
ders Ranges, South Australia. This species from Pararaia
13
G. Geyer et al.
composite mould illustrating differences in reticulation on the shell
exterior and interior, sample locality W8; ventral view; k, l, MMUW
2014A-113, latex cast of external mould in h, with very prominent
reticulate ornamentation of the shell exterior, sample locality W8;
oblique lateral and dorsal views; m, MMUW 2014A-168a, partial
conch, detail of shell exterior with prominent reticulate riblets, sam‑
ple locality W8. All specimens from Tannenknock Formation, Wil‑
denstein Member. All scale bars 1  mm. N, Helcionella lemdadensis
sp. nov., SMF 34737, Lemdad syncline, High Atlas, Morocco, section
Le II, sample horizon X255 (specimen figured in Geyer, 1986, pl. 1,
fig. 9). Scale bar 1 mm
bunyerooensis Zone (Cambrian Stage 3) is distinguished
from H. lemdadensis (and H. oblonga) in having a coarser
and more prominent reticulation and more sharply developed
concentric ribs.
Helcionella aff. oblonga Cobbold, 1921
Figure 5a–c, e–i, k–m, d?, j?
Material. 20–25 specimens, mostly internal moulds and
composite moulds. From sample locality W8: MMUW
2014A-105, MMUW 2014A-111a, MMUW 2014A-111b,
MMUW 2014A-113, MMUW 2014A-146a, MMUW
2014A-147a, MMUW 2014A-148a, MMUW 2014A-148b,
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
MMUW 2014A-148c. From sample locality W8: MMUW
2014A-127, MMUW 2014A-128, MMUW 2014A-131,
MMUW 2014A-132, MMUW 2014A-133, MMUW 2014A-
134, MMUW 2014A-135a, MMUW 2014A-168a, MMUW
2014A-173, MMUW 2014A-174a, SSMM 11355, SSMM
10795. Tentatively assigned to H. aff. oblonga: From sam‑
ple locality W8: MMUW 2014A-107, MMUW 2014A-129,
MMUW 2014A-149a, MMUW 2014A-152, SSMM 10408,
SSMM 11351, SSMM 11357. From sample locality W8:
MMUW 2014A-125a.
Localities and stratum. Wildenstein slice, locality W8,
Franconian Forest. Tannenknock Formation, Wildenstein
Member.
Description. Isostrophic shell forms a cap-shaped, com‑
pressed cyrtoconic conch with a large suboval to slightly
subrectangular aperture. Maximum length of the present
specimens ranges from ca. 3 to 8  mm, with an average
width/length ratio of ca. 0.75. In the early growth stages, the
cross-section is suboval, and the costae extend around the
entire shell in subequal breadth, but they gradually become
narrower along the sub-apical area so that their margins
describe more and more obvious acute angles in lateral
view and become increasingly prominent as the aperture is
approached. Internal moulds are devoid of pseudoseptate
incisions.
Shell exterior with finely reticulate pattern on the conch;
interior with similar, but very fine reticulate pattern.
Discussion. The specimens from the Franconian Forest are
similar to Helcionella capula from the same formation, but
differ in the absence of pseudoseptate incisions, a more
strongly extended apical portion, more prominent, rounded
rugae, and a generally larger size. They match in morphol‑
ogy with Helcionella oblonga Cobbold, 1921 from the
Paradoxides groomi Grits (now the basal part of the Upper
Comley Sandstone) of Comley, Shropshire, Britain, and a
specimen subsequently found in the Lemdad syncline, High
Atlas, Morocco (Geyer 1986). However, differences from
these can be seen in a more strongly protruding apical por‑
tion, a less rapid growth of the aperture and a comparatively
more slender outline. Nevertheless, the preservation of the
specimens from the Franconian Forest does not allow a con‑
fident identification.
Helcionella sp. A
Figure 6a–o, q–s, p?
Material. Ca. 20 specimens, mostly internal moulds and
composite moulds. From sample locality W12 (Galgenberg
Member): MMUW 2014A-101a. From sample horizon
W8 (Wildenstein Member): MMUW 2014A-103, MMUW
217
2014A-163a, MMUW 2014A-169Ia and 2014A-169IIa,
MMUW 2014A-170a, MMUW 2014A-177, MMUW
2014A-178a, MMUW 2014A-181, MMUW 2014A-182,
MMUW 2014A-183a, MMUW 2014A-185a, MMUW
2014A-186a, SSMM 10407, SSMM 10799, SSMM 10829a,
b (counterparts), SSMM 11077, SSMM 11354. Tentatively
assigned to Helcionella sp. A: From sample locality W15a
(Wildenstein Member): MMUW 2014A-102. From sample
locality W8: MMUW 2014A-150, MMUW 2014A-189a,
MMUW 2014A-189b, MMUW 2014A-190. From sample
locality W8: MMUW 2014A-123a.
Localities and stratum. Wildenstein slice, Franconian For‑
est. Tannenknock Formation, Galgenberg and Wildenstein
members.
Description. Isostrophic shell forms a cap-shaped, moder‑
ately expanded to somewhat compressed, slightly cyrtoconic
conch with a large suboval to faintly subrectangular aperture.
Maximum length of the present specimens ranges from ca.
5 to 11 mm, with an average width/length ratio of ca. 0.80.
The plications extend around the entire shell in subequal
breadth in the juvenile part of the conch, but they become
somewhat more irregular towards the apertural margin.
Shell exterior with distinct concentric incisions which are
relatively shallow on the supra-apical surface of the shell,
but more distinctly incised on the sub-apical surface under‑
neath the apex, there resembling the distinct incisions seen
in Helcionella capula (see previous). Ribs between the con‑
centric incisions usually with low convexity in lateral view,
covered by densely spaced, moderately coarse subparallel
riblets arranged perpendicular to the incisions (Fig. 6q–s).
These riblets are generally aligned in such a way to be paral‑
lel to the riblets of the adjacent plications and thus would
be seen as almost straight lines stretching from the apical
part to the apertural margin if not dissected by the incisions
(Fig. 6l). Thus, the riblets grow slightly in width towards the
apertural margin (Fig. 6q). Rare intercalations of new riblets
can be seen in the plications close to the apertural margin.
The shell interior has a similar, fine reticulate pattern, which
is rarely preserved in the present specimens (Fig. 6k).
Discussion. The specimens from the Franconian Forest
treated here as Helcionella sp. A are similar to the co-occur‑
ring Helcionella aff. oblonga Cobbold, 1921, and incomplete
specimens with unfavourable preservation may be confused
with them. However, Helcionella sp. A is unequivocally
distinguished by a low apical portion that creates an only
slightly cyrtoconic shape of the conch so that the specimens
are often preserved as low specimens with a suboval outline
in a strongly dorsoventrally flattened condition.
The species represented by the specimens also resembles
Helcionella capula Geyer, 1986, but differ in the absence of
13

218 G. Geyer et al.

13
 Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
◂Fig. 6  a–o, q–s, p? Helcionella sp. A. a, b SSMM 10829a, b, large
conch, Tannenknock Formation, Wildenstein Member, sample local‑
ity W8; a internal mould, oblique view; B, external mould, oblique
view, counterpart of specimen in a; c, f SSMM 10407, small conch,
laterally compressed; Tannenknock Formation, Wildenstein Mem‑
ber, sample locality W8; dorsal and lateral views; d, j, n MMUW
2014A-169aI, small conch, internal mould, dorsal, lateral and pos‑
terior views; Tannenknock Formation, Wildenstein Member, sam‑
ple locality W8; e MMUW 2014A-101a, dorsoventrally compressed
conch typical for Galgenberg Member of the Tannenknock For‑
mation, sample horizon W12?; oblique lateral view; g, k MMUW
2014A-111b, latex cast of external mould, sample locality W8; lateral
and dorsal views; h MMUW 2014A-103, internal mould, Tannen‑
knock Formation, Wildenstein Member, sample horizon W8; oblique
lateral view; i, l, m MMUW 2014A-181, dorsoventrally flattened and
crushed specimen with exposed apical internal mould; Tannenknock
Formation, Wildenstein Member, sample locality W8; i oblique ante‑
rior view; l dorsal view; m detail of external surface in dorsal view; o
MMUW 2014A-182, external mould of partial conch, ventral view;
Tannenknock Formation, Wildenstein Member, sample locality W8;
p MMUW 2014A-123a, partial conch, example for poorly preserved
specimen; Tannenknock Formation, Wildenstein Member, sample
locality W8; q, r, s MMUW 2014A-163a, conch with preserved,
fractured shell, dorsal, lateral and posterior views; Tannenknock For‑
mation, Wildenstein Member, sample locality W8. Scale bars 1 mm.
Photos a, b courtesy S. Meier, Marktredwitz
clear “pseudoseptate” incisions, a more subcentral position
of the apex, a generally larger size, and probably a more
strongly developed reticulation of the shell exterior.
The preservation of the specimens from the Franconian
Forest is not sufficiently favourably to allow a precise char‑
acterization of the species. A number of Helcionella spe‑
cies resemble the specimens from the Franconian Forest,
particularly H. subrugosa (d’Orbigny, 1850) from the late
early Cambrian of the New York State, but most are clearly
distinguished in having the apex in a less subcentral position
and being more elevated (despite of the dorsoventral flatten‑
ing of the Franconian specimens).
Genus Bemella Missarzhevsky in Rozanov et al., 1969
Type species. Helcionella jacutica Missarzhevsky in Rozanov
and Missarzhevsky (1966), from the lower Cambrian Doki-
docyathus regularis Zone, Tommotian Stage, middle part of
Ken River, Lena-Aldan region, Siberian Platform.
Discussion. The generic concept of Bemella has been
modified several times due to an inadequately precise dis‑
crimination in the first studies from the Siberian Platform.
Subsequent findings of Bemella-type specimens from other
regions and younger strata than those yielding the Sibe‑
rian specimens revealed deficiencies in the initial generic
concept. The subsequent emendation by Parkhaev (2001)
restricted Bemella to cap-shaped, slightly coiled cyrto‑
conic helcionelloids with a somewhat laterally compressed
conch covered by comarginal rugae and with a simple, plan
219
aperture. This, in fact, allows for the occurrence of tran‑
sitional stages between Helcionella and Ilsanella (as dealt
with by Parkhaev 2001), and thereby invalidates the mor‑
phological significance of the genera. As noted before, an
alternative approach to the generic concepts by Jacquet and
Brock (2016) distinguished the genera by the height of the
conchs, rate of expansion and non-bifurcating rugae, but
appears to be an attempt select any possible characters that
allow a more or less confident unequivocal distribution of
the species between the genera.
It must be emphasized that the large number of species
attributed to Bemella (at least 19 species in Parkhaev 2001
plus a few others not cited therein) partly exhibit differences
that may be regarded as of supraspecific significance (such
as distinct differences in the profiles of the ribs, the shape
of the sub-apical areas, and the morphology and definition
of the larval part of the shell) so that the genus appears to
represent a polyphyletic unit (e.g., compare B. jacutica (Mis‑
sarzhevsky in Rozanov and Missarzhevsky, 1966) with B.
incomparabilis Parkhaev, 2001, and B. wiri Kruse, 1998).
A revision of the genus is urgently needed but beyond the
scope of this study.
“Bemella” sp. A
Figure 7e, f, h, j
Material. Two (possibly three) specimens. From W8:
MMUW 2014A-119, MMUW 2014A-121; tentatively
assigned to “Bemella” sp. A from the same locality and
horizon: MMUW 2014A-188. All from Wildenstein slice,
Franconian Forest, Tannenknock Formation, Wildenstein
Member.
Description. Low cyrtoconic, quasi cap-shaped shell, mod‑
erately low, with plan, oval aperture. Apex blunt, inclined
posteriorly/sub-apically, located approximately at rear edge
of the aperture in dorsal view. Supra-apical surface gently
convex in the apical area, considerably flattened towards
the aperture in lateral view. Sub-apical surface slightly con‑
cave. Lateral fields faintly convex in the abapical part of
the shell. Shell exterior covered by relatively coarse, more
or less regular concentric ribs, which are clearly broader
across supra-apical surface, diminished towards sub-apical
surface. Larger of the present specimens with probably 7
ribs. Corrugation smoothing toward apical region. Apical
part (protoconch?) appears to be an oval, faintly bulbous
part, sitting slightly oblique on the teleoconch.
Discussion. The two specimens found in the Tannenknock
Formation are too poorly preserved to allow a morphologi‑
cally sound description of a species. The morphology resem‑
bles several species described under Bemella, particularly
in the relatively blunt and dorsoventrally elongate apex.
13

220 G. Geyer et al.

Fig. 7  a–d, g, i, k, m Helcionel‑
loid gen. and spec. indet. A.
a, b, g MMUW 2014A-144,
incomplete internal mould,
sample locality W13b; lateral,
dorsal and posterior views; c, d
SSMM 11551, small conch with
partly preserved shell; i, k, m
MMUW 2014A-114, incom‑
plete internal mould, sample
locality W8; anterior and lateral
views. e, f, h, j “Bemella” sp.
A. e, f MMUW 2014A-119,
incomplete internal mould,
sample locality W8; lateral and
posterior views; h, j MMUW
2014A-121, incomplete internal
mould, sample locality W8;
lateral and dorsal views. l
Yochelcionella? sp. indet.,
SSMM 11601, incomplete
internal mould with anteriorly
directed minute apical portion
(ap) and snorkel-type tube
(sn), sample locality W8. All
scale bars 1 mm. All specimens
from Tannenknock Formation,
Wildenstein Member. Photo L
courtesy S. Meier, Marktredwitz

13
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
However, the conchs are less laterally compressed than most
of the species under the generic concept of Parkhaev (2001).
The specimens from the Franconian Forest bear close resem‑
blance to Bemella bella Chen and Zhang, 1980 from the late
Terreneuvian of Yunnan, South China Platform. However,
the similarity is regarded as based on plesiomorphic charac‑
ters and is insufficient to prove a close relationship between
them. It is this combination that prompted the recognition
of B. bella in Avalonian eastern Massachusetts and south‑
eastern Newfoundland as well (Landing 1988; Landing et al.
1989).
Genus Igorella Missarzhevsky in Rozanov et al., 1969
Type species. Igorella ungulata Missarzhevsky in Rozanov
et al. (1969), from the lower Cambrian Tommotian Stage,
West Anabar and Uchur-Maya regions, Siberian Platform.
Discussion. The morphological concept of Igorella awaits
a clarification after the genus has been used increasingly to
provide somewhat laterally compressed helcionellid-type
conchs with a plan aperture and an apical portion overhang‑
ing the sub-apical field. Lateral compression, as well as
the position of the apex and the external ornament of the
shell varies considerably between the species assigned to
the genus (e.g. I. ungulata Missarzhevsky in Rozanov et al.
1969, pl. IV, figs. 12, 15, 21; Igorella emeiensis (Yu, 1987)
in Parkhaev 2005a, pl. II, fig. 1a, b; and Igorella maidipin-
gensis (Yu, 1974) in Parkhaev 2005a, pl. II, fig. 2a, b). The
genus is here understood as being characterized largely by
its type species, I. ungulata Missarzhevsky in Rozanov et al.
(1969), the aperture of which has a slender elliptical shape,
the apex being moderately broad in lateral profile on the
exterior of the shell and relatively acute on internal moulds,
and the exterior of the shell with fairly low, densely spaced
and somewhat irregular comarginal rugae. Similar forms are
almost globally distributed (e.g., I. durara Kruse, 1991 from
the lowermost middle Cambrian Top Springs Limestone of
the Georgina Basin, Australia).
“Igorella” sp. A
Figure 8
Material. Four specimens. From sample locality S1: SSMM
11557, SSMM 11558a, and SSMM 11559; from W8:
MMUW 2014A-191c. Schnebes–Wustuben and Wildenstein
slices, Franconian Forest; Tannenknock Formation, Wilden‑
stein Member, and lower part of the Triebenreuth Formation.
Description and discussion. The material consists of internal
moulds of a cyrtoconic shell, which are moderately low and
possess a plan, oval aperture with a width/length ratio of ca.
0.5 to 0.6. The conch is considerably compressed laterally
221
in the juvenile part, but expands moderately toward the
aperture. The apex is small, slightly compressed laterally
and inclined posteriorly/sub-apically, and it appears to be
quite well defined from teleoconch as indicated by specimen
MMUW 2014A-191c (Fig. 8f). It is located approximately at
or near the rear edge of the aperture in dorsal view. The sub-
apical surface is concave in lateral view and has a narrow
curvature in transverse section. The lateral fields are faintly
to moderately convex and lack distinct ribs or corrugations.
The single specimen of “Igorella” sp. A recovered from
Wildenstein Member of sample locality W8 is supplemented
by several, better preserved specimens from the Triebenreuth
Formation so that they are included herein to illustrate its
morphology.
Helcionelloid genus and species indeterminate A
Figure 7a, b, e, g, i, k
Material. Four specimens. From sample locality W8:
MMUW 2014A-114, SSMM 10791a, SSMM 10792; from
sample locality W13b: MMUW 2014A-144. Wildenstein
slice, Franconian Forest; Tannenknock Formation, Wilden‑
stein Member.
Description. Cyrtoconic, cap-shaped shell, moderately
low, with plan, oval aperture. Apex recurved posteriorly,
located approximately at rear edge of the aperture in dor‑
sal view. Shell exterior covered by moderately coarse, more
or less regular concentric ribs (up to seven in the present
specimens), which are broader across supra-apical surface,
diminished towards sub-apical surface. Corrugation smooth‑
ing toward apical region. Apical part appears to be slightly
oval, but subacute, without recognizable differences to tel‑
eoconch. A better preserved of the specimen from the lower
part of the Triebenreuth Formation near Wustuben (SSMM
11551; Fig. 7c) has the shell retained for most part of the
conch, which indicates that the external surface was smooth.
Discussion. The specimens found in the Tannenknock For‑
mation are imperfectly preserved and do not allow recog‑
nition of significant characters. In addition, they are small
and appear to represent shells of juvenile individuals. The
morphology bears resemblance to several species described
under Bemella, but the conchs are less laterally compressed,
have a subacute rather than blunt apex with an oval cross-
section and possess comparatively narrow and more tightly
spaced concentric ribs.
Family Coreospiridae Knight, 1947
Remarks. A parietal train that causes a groove or indenta‑
tion at the posterior/sub-apical apertural margin has been
regarded by Parkhaev (2001, 2008) as a diagnostic character
13

222 G. Geyer et al.

Fig. 8  “Igorella” sp. A. a, b,
SSMM 11557, internal mould
of conch, lateral and dorsal
views, from sample horizon S1;
c, d, SSMM 11558a, internal
mould of conch, oblique lateral
and dorsal views, from sample
horizon S1; e, f, MMUW
2014A-191c, internal mould of
conch, dorsal and lateral views,
from sample horizon W8. All
scale bars 1 mm (equivalent to
magnification ×20). a–d From
Triebenreuth Formation, near
Wustuben, Schnebes–Wustuben
slice; e, f from from Tannen‑
knock Formation, Wildenstein
Member, Wildenstein slice

that separates the superfamily Yochelcionelloidea sensu
Parkhaev (composed of the families Yochelcionellidae,
Trenellidae and Stenothecidae) from the superfamily Hel‑
cionelloidea (Helcionellidae, Coreosiridae and Carinopelti‑
dae [= Igarkiellidae]) which lacks a parietal train (Parkhaev,
2001, 2008, 2017). Therefore, this feature also distinguishes
the members of the family Trenellidae Parkhaev, 2001 from
typical coreospirid genera such as Latouchella Cobbold,
1921. However, the development and expression of a pari‑
etal train appears to be transitional in a number of forms
and can only be scrutinized for the taxon by well-preserved
specimens, whereas a close relationship and evolutionary
derivation of genera such as Latouchella and Oelandia
Westergård, 1936 appears to be obvious. However, Parkhaev
(2001, 2008) interpreted Latouchella as from a lineage of
planispirally coiled species typical for the Coreospiridae,
whereas he reconstructs Oelandia as an uncoiled train-bear‑
ing genus which not only affects the validity of a separation
into the families Coreospiridae and Trenellidae, but also that
of the superfamilies Helcionelloides and Yochelcionelloidea
in Parkhaev’s concept.
Genus Latouchella Cobbold, 1921
Type species. Latouchella costata Cobbold, 1921, from the
Lower Comley Limestone of Shropshire, Great Britain.
Latouchella cf. comma Geyer, 1986
Figure 9
Material Ca. one dozen specimens. From sample local‑
ity W8: MMUW 2014A-115, MMUW 2014A-116, and

13
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany 223

Fig. 9  Latouchella cf. comma Geyer, 1986. a MMUW 2014A-100, ▸

incomplete conch, internal mould, obliquely compressed, Tannen‑
knock Formation, Wildenstein Member, sample locality W13a, lateral
view; b, d, g MMUW 2014A-115, fragment of conch, internal mould,
with differently developed ribs, Tannenknock Formation, Wildenstein
Member, sample locality W8, lateral views and view perpendicular to
apertural plane; c, e MMUW 2014A-158a, latex cast of incomplete
conch, internal mould, Tannenknock Formation, Wildenstein Mem‑
ber, sample locality W8, lateral and oblique views; f MMUW 2014A-
116, fragment of conch showing exterior of shell with faint oblique
striae, Tannenknock Formation, Wildenstein Member, sample local‑
ity W8; h–k MMUW 2014A-162, fragment of conch, internal mould
with prominent ribs, Tannenknock Formation, Wildenstein Member,
sample locality W8; different views, k shows lateral view with faint
striae perpendicular to the longitudinal axes of the ribs; l MMUW
2014A-187, incomplete conch, external mould, Tannenknock For‑
mation, Wildenstein Member, sample locality W8; m, n MMUW
2014A-184, incomplete conch, external mould, Tannenknock Forma‑
tion, Wildenstein Member, sample locality W8. Scale bars 1 mm

MMUW 2014A-158a. From W8: MMUW 2014A-162,

MMUW 2014A-184, MMUW 2014A-184, MMUW 2014A-
187, SSMM 10422, SSMM 10815, SSMM 10816, and
SSMM 10817. Tentatively assigned to L. cf. comma: from
sample locality W13a: MMUW 2014A-100.

Localities and stratum. Wildenstein slice, Franconian Forest,

localities W8 and W13a. Tannenknock Formation, Wilden‑
stein Member.

Description. The specimens from the Franconian Forest are

the remains of a small univalved mollusc with a maximum
length of about 5 mm and a maximum width of about 2 mm,
represented by internal moulds coiled through approximately
one whorl. The apical portion of the shell is not preserved
in any of the specimens. The aperture is oval and elongate
(length/width ratio ca. 2:1). The lateral surface of the whorl
has slender, but prominent ribs which disappear towards the
sub-apical and the dorsal surfaces so that they do not cross
the dorsum. A specimen assigned to the same form shows a
pattern of faint growth lines in an acute angle with the ribs
and extremely faint lirae (Fig. 9f).

Discussion. The available fragmentary material is insuffi‑

cient to allow a more precise determination. However, the
specimens match Latouchella comma Geyer, 1986 in all
aspects. This species is known from the lower part of the
middle Cambrian in the High Atlas Mountains, Morocco,
and occurs there in strata that are perfectly coeval.
As noted by Geyer (1986, p. 79) Latouchella costata Cob‑
bold, 1921, the type species of the genus from the Lower
Comley Limestone of Shropshire, is differentiated from L.
comma by slightly stronger coiling. In addition, the lateral
ribs in L. costata are less extended towards the dorsum. The
otherwise similar Latouchella penecyrano Runnegar and

13

224 G. Geyer et al.

Jell, 1976 is differentiated by its less prominent ribs and a
more slender transverse section.
Genus Leptostega Geyer, 1986
Type species. Leptostega irregularis Geyer, 1986, from early
middle Cambrian strata of the Láncara Formation, Canta‑
brian Mountains, Spain.
Emended diagnosis. Genus of the Coreospiridae with fol‑
lowing characters: conch narrow, subtriangular in lateral
view, with an apex dipping gently in posterior direction;
lateral sides weakly convex; surface traversed by concen‑
tric ribs, which alternate or are connected in an irregular
arrangement at the narrow anterior and posterior sides.
Remarks. The concept of Leptostega as introduced by Geyer
(1986) has been poorly understood, probably as a result of the

13
 Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
◂Fig. 10  a–t Leptostega frankenwaldensis sp. nov. a, b MMUW
2014A-139, paratype, incomplete conch, composite mould with faint
striae, Tannenknock Formation, Wildenstein Member, sample local‑
ity W18a, lateral and oblique posterior views; c, d, i MMUW 2014A-
138, holotype, incomplete conch, internal mould, Tannenknock For‑
mation, Wildenstein Member, sample locality W18a, lateral, apical
and anterior views; e, j MMUW 2014A-136a, paratype, partial conch,
internal mould, Tannenknock Formation, Wildenstein Member, sam‑
ple locality W8, lateral and apical views; f MMUW 2014A-153a,
paratype, incomplete conch, internal mould, Tannenknock Formation,
Wildenstein Member, sample locality W8, lateral view; g, h MMUW
2014A-140a, paratype, incomplete conch, composite mould with faint
striae, Tannenknock Formation, Wildenstein Member, sample local‑
ity W18a, lateral and apical views; k MMUW 2014A-154, paratype,
partial conch, composite mould, Tannenknock Formation, Wilden‑
stein Member, sample locality W8, lateral view; l, m, n MMUW
2014A-161a, paratype, incomplete conch, internal mould, Tannen‑
knock Formation, Wildenstein Member, sample locality W8, apical,
oblique lateral and lateral views; o SSMM 10419b, paratype, incom‑
plete conch, internal mould, Tannenknock Formation, Wildenstein
Member, sample locality W8, lateral view; p MMUW 2014A-164a,
paratype, conch, internal mould, Tannenknock Formation, Wilden‑
stein Member, sample locality W8, lateral view; q, r, s, t MMUW
2014A-161b, paratype, incomplete conch, internal mould, Tannen‑
knock Formation, Wildenstein Member, sample locality W8, lateral,
anterior and posterior views. u–y Helcionelloid genus and species B.
u–x MMUW 2014A-192, latex cast of external mould of conch, lat‑
eral, oblique anterior, and oblique apical views, showing lateral ribs
extending with reduced height around posterior face and absence of
ribs on anterior face; note presence of narrowly spaced growth lines
(arrow in v); y MMUW 2014A-193, latex cast of external mould, lat‑
eral view showing oblique course of lateral ribs and its reduced width
towards posterior face, with fine growth lines visible between adap‑
ertural ribs and on adapertural rib (left side). Both specimens from
Tannenknock Formation, Wildenstein Member, sample locality W8.
All scale bars 1 mm
language in which that article has been published. Essential
characters are the laterally compressed conch and the sub‑
central apex with only a gentle posterior inclination. Mor‑
phologically similar is Mackinnonia/Davidonia Parkhaev,
2017b (Davidonia replaces Mackinnonia Runnegar in Bengt‑
son et al., 1990 according to Parkhaev 2017b; see discussion
below), which has a moderately compressed conch and a pos‑
teriorly displaced, hook-shaped apical portion in a cap-shaped
shell. Oelandia Westergård, 1936 is also closely related to
Leptostega and Mackinnonia/Davidonia. Oelandia is charac‑
terized by an alternation of the coarse comarginal plications
from one side of the shell to the other, producing a bilateral
asymmetry (see Peel and Yochelson 1987, for details). This
alternation is similar to that seen in specimens of Leptostega
irregularis, and also produces faint processes on the abapical
side. Tannuella Missarzhevsky, 1969 has a subcentral apex,
but a less compressed conch and coarse concentric ribs.
Parkhaev (in Gravestock et al. 2001) placed Isitella plicata
Missarzhevsky, 1989 under Mackinnonia, which is regarded
here as correct (in case Mackinnonia is not validly replaced
by Davidonia). Mackinnonia plicata, however, has concentric
prominent ribs with rounded rectangular profiles, which often
225
show distinct arched folds towards the apical portion on the
supra-apical surface. The same features are seen in Leptostega
irregularis described below.
Parkhaev (2017b) discussed that Mackinnonia Runnegar,
1990 is a junior homonym of Mackinnonia Janiszewska,
1963 (a Recent myriosporid sporozoan; type species: M.
tubificis Janiszewska, 1963) and suggested the replace‑
ment name Davidonia. However, Jackson and Claybourn
(2018) emphasize that Mackinnonia Janiszewska, 1963
must be regarded as a nomen nudum: Janiszewska (1963)
did not provide a diagnosis of Mackinnonia, did not desig‑
nate formally a type species or type material or any specific
characterization, and a sometimes purported earlier use of
name in Janiszewska (1957) does not exist. Consequently,
Mackinnonia Runnegar, 1990 claims priority, and Davidonia
Parkhaev, 2017 must be regarded as a junior synonym of the
latter genus. Nevertheless, Mackinnonia Janiszewska, 1963
is in use, M. tubificis being described and figured in some
detail in Janiszewska (1967) (albeit obviously without mate‑
rial being deposited), and an additional species (M. lumbri-
cilli Siau and Ormières, 1970) is established.
Landing (1988) redescribed and figured the type of Sten-
otheca rugosa var. abrupta Shaler and Foerste, 1888 from
the Callavia Zone of the Weymouth Formation of eastern
Massachusetts, which he designated as Helcionella abrupta.
This species shows a similar pattern of thickened folds with
an apical curvature on the anterior side and an overall com‑
prehensive similarity with Mackinnonia plicata, which
favours a placement under Makinnonia. However, accord‑
ing to the single specimen known thus far, the species from
western Avalonia appears to differ in inserted ribs and a
smaller apex so that Parkhaev’s (2001) synonymization with
M. plicata appears to be premature until better preserved
material is available from the Weymouth Formation. For the
moment, the species should be dealt with as Mackinnonia?
abrupta. An incomplete specimen very reminiscent of the
type of M.? abrupta has been described and figured from
the upper lower Cambrian Forteau Formation of Laurentian
western Newfoundland as Mackinnonia sp. by Skovsted and
Peel (2007, fig. 4b).
Mackinnonia puppis Høyberget, Ebbestad, Funke et
Nakrem, 2015 from the uppermost lower Cambrian Evjevik
Member of the Ringstrand Formation in the Møsa District,
Norway, resembles Leptostega, particularly in the nodular
thickenings on sub-apical area as in L. frankenwaldensis sp.
nov. The species from southern Norway, however, is much
larger and has the concentric plicae typical of Mackinnonia.
Leptostega frankenwaldensis sp. nov.
Figure 10a–t
Etymology. Named after its occurrence in the Franconian
Forest range (German: Frankenwald).
13

226
Holotype. MMUW 2014A-138, fairly complete internal
mould of conch (Fig. 10c, d, i).
Type locality and type stratum. Sample locality W18a north
of Wildenstein, Franconian Forest; Tannenknock Formation,
Wildenstein Member.
Paratypes. 19 specimens, mostly internal moulds. From
sample locality W18a: MMUW 2014A-139, MMUW
2014A-140a. From sample locality W8: MMUW
2014A-136a, MMUW 2014A-153a, MMUW 2014A-154,
MMUW 2014A-161a, MMUW 2014A-161b, MMUW
2014A-164a, MMUW 2014A-166a, MMUW 2014A-172a,
MMUW 2014A-176, SSMM 10419b, SSMM10790, SSMM
11036e, and SSMM 11198b.
Localities and stratum. Wildenstein slice, Franconian Forest,
sample localities W8 and W18a; Tannenknock Formation,
Wildenstein Member.
Diagnosis. Species with slightly cyrtoconic shell with an
aperture having a narrow elliptical outline with a length/
width ratio of ca. 1.4 to 1.8. Ribs with progressively nodular
thickenings on sub-apical area. Apex located at ca. 35 to 38
percent maximum length.
Description. Shell distinctly compressed laterally, with gen‑
tly convex sides, more strongly expanding terminal abapical
portion, subtriangular to slightly cyrtoconic in lateral view.
Aperture with narrow elliptical outline; length ca. 1.4‒1.8
times greater than its width. Apex small, barely bulbous,
slightly displaced anteriorly, gently oblique to vertical axis.
Supra-apical surface of the shell very gently and almost
evenly convex except for sinuosities created by ribs; ribs
slightly swollen in breadth by comparison to its width on
lateral fields (e.g., Fig. 10c). Sub-apical surface of the shell
more or less straight in total, but ribs continuing. Internal
moulds with 4–5 broad concentric ribs with sinuous profile
on the lateral sides, width of ribs and grooves roughly equal,
but subject to considerable variation. Ribs of the lateral sides
pass into distinctly thickened folds with an apical curva‑
ture on the posterior side so that the ribs there are distinctly
broader than the grooves between them; a similar pattern of
thickened ribs can be seen on the posterior/sub-apical side
that may create low thorn-like protuberances (Fig. 10c, f, q).
Exterior of shell not well preserved in any of the speci‑
mens, internal surface sometimes with faint striae arranged
perpendicular to concentric ribs (e.g., Fig. 10g, p).
Remarks. The specimens from the Wildenstein Formation
are similar morphologically to the type material of Lep-
tostega irregularis Geyer, 1986 from the almost coeval strata
of the Láncara Formation in the Cantabrian Mountains,
13
G. Geyer et al.
Spain. Differences exist in the broader and clearly less prom‑
inent ribs of the specimens from the Wildenstein Formation;
the more regular ribbing of the conchs; and the more upright
direction of the apex.
Leptostega cingulata (Cobbold, 1921) is known only
from a single, small (ca. 2 mm) fragment from the Protole-
nus Limestone of the Lower Comley Limestone of Comley,
Shropshire, which is characterized by more numerous (at
least 7), narrow, prominent ribs without recognizable swell‑
ings on the sub-apical surface.
An additional species has been attributed to Leptostega as
L. hyperborea Parkhaev, 2005b. This species from the upper
part of the Emyaksin Formation, Botoman Stage, of the Ana‑
bar River basin, Yakutia, nicely portrays, in comparison to
Leptostega cingulata, the other extreme morphology. It has
a conch with a superficial, almost bilaterally symmetrical
outline in lateral view, in which the few (four), thick and low
ribs on the lateral sides proceed into conspicuously swol‑
len processes on the narrow, apical and abapical fields, and
these are separated by incisions similar to the pegma-type
sinuosities (Parkhaev 2005b, pl. 2, fig. 1a–f). A similar form
has been described by Kouchinsky et al. (2011) from the late
Amgan of the upper Kuonamka Formation, Bol’shaya Kuon‑
amka River section, Siberian Platform, as Leptostega sp. cf.
L. hyperborea. Specimens representing this form are incom‑
plete internal moulds. The ribs are more slender than in the
distinctly older L. hyperborea, which is in part attributable to
the preservation as internal moulds. However, the processes
on the narrow sub-apical field are less pronounced, and the
cross-section has a more lenticular shape than L. hyperbo-
rea, therefore it most probably represents a new species of
Leptostega. It should be emphasized that their outline in
lateral view appears to vary considerably as demonstrated by
the specimens in Kouchinsky et al. (2011, fig. 7a, e).
A species from the lower Cambrian Parabadiella
(“Abadiella”) huoi Zone of the Parara and Ajax Lime‑
stones as well as the Oraparinna Shale of South Australia
has been tentatively assigned to the genus as Leptostega?
corrugata Runnegar, 1990 (in Bengtson et al. 1990, p. 234,
fig. 160a–g). The species has been restudied by Parkhaev
(2001) and synonymized with Isitella plicata Missarzhevsky,
1989, which Parkhaev (for reasons that were not explicitly
discussed) placed among the genus Mackinnonia Runnegar
in Bengtson et al., 1990 (now Davidonia sensu Parkhaev,
2017b). The position under Mackinnonia/Davidonia is obvi‑
ous, but we do not follow the synonymization proposed by
Parkhaev (2001) as already discussed by Skovsted (2004).
As pointed out by Parkhaev (2001, 2005a) Leptostega? cor-
rugata shows the distinctly cyrtoconic shape of the conch
rather than the slightly inclined apical portion as in L. irreg-
ularis, and a regular plication, which, however, culminates
in the very prominent, sometimes protuberance-type ribs on
the abapical and sub-apical surface so that its morphological
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
characters agree better with Mackinnonia rather than Lep-
tostega. It should be noted, however, that one specimen
figured by Gravestock et al. (2001, pl. XXXIX, fig. 7) has
a different morphology than the other specimens assigned
to Mackinnonia plicata and resembles the specimens intro‑
duced here as Leptostega frankenwaldensis sp. nov.
In addition, Parkhaev (2001) also synonymized Sten-
otheca rugosa var. abrupta Shaler and Foerste, 1888 with
Isitella plicata under Mackinnonia plicata. This species is
known only from a poorly preserved specimen from the
lower Cambrian Callavia Zone/Serrodiscus bellimargina-
tus fauna of the Weymouth Formation in Avalonian eastern
Massachusetts, best refigured in Landing (1988, fig. 5.14;
as Helcionella abrupta). Another specimen attributed to
the species as Stenotheca abrupta by Grabau (1900, p1.
XXXI, fig. 12a–c) certainly belongs to a different species.
The species shows a notable swelling of its ribs towards
the narrow sides, but the crests of these ribs are distinctly
corroded so that the true nature cannot be restored. The sug‑
gested synonymy is difficult to demonstrate and verify and at
least highly uncertain. However, if the synonymy would be
correct, the species would have to be named Mackinnonia
abrupta (Shaler and Foerste, 1888) rather than M. plicata
(Missarzhevsky, 1989).
Helcionelloid genus and species B
Figure 10u–y
Material. Two specimens, external moulds. From sample
locality W8: MMUW 2014A-192, MMUW 2014A-193.
Wildenstein slice, Franconian Forest, Tannenknock Forma‑
tion, Wildenstein Member.
Description. Cyrtoconic shell, laterally compressed, with
elongate elliptical transverse section. Lateral sides gently
convex. Aperture with elliptical outline, length roughly 1.6
times greater than width, planar for most part, but with slight
apicalward curvature at the sub-apical end. Apical portion
directed slightly posteriorly, apex small, located approxi‑
mately posterior slightly to centre of shell. Supra-apical sur‑
face of the shell gently and almost evenly convex. Sub-apical
surface with slight concave curvature in lateral view, with
low ribs. Shell with at least seven concentric ribs, which are
best developed and highest on the lateral sides, where they
are almost straight, but somewhat inclined toward posterior,
extending as low ribs onto the sub-apical surface; width of
ribs greater than that of concavities between them in the
apical part, subequal in the middle part of the conch and
slightly narrower in the adaxial portion. Surface with fine,
narrowly spaced growth lines (Fig. 10u, x).
Discussion. The form represented by the two conchs
are characterized by its laterally compressed shells with
227
numerous straight, low ribs and a triangular outline in lateral
view that resemble species of the genus Stenotheca Salter
in Hicks (1872). However, the form is distinguished by its
nearly planar aperture without a distinct parietal train, by
a more broadly elliptical transverse profile, by the absence
of ribs on the supra-apical surface, and by the subcentral
position of the apex. The form belongs to the family Core‑
ospiridae rather than the family Stenothecidae according to
the generally accepted systematic concept of Coreospiridae.
Nevertheless, it appears to be obvious that Stenotheca with
its slightly curved aperture may be derived from this type
of helcionelloid, reflecting the development from an animal
moving on the sediment surface (or slightly plowing through
the topmost millimetres if soft mud) into a semi-infaunal
mode of lifen a suggested for the Stenothecidae.
Family Yochelcionellidae Runnegar and Jell, 1976
Genus Yochelcionella Runnegar and Pojeta, 1974
Type species. Yochelcionella cyrano Runnegar and Pojeta,
1974 (by original designation); Lower Cambrian Coonigan
Formation, New South Wales, Australia.
Yochelcionella? sp. indet.
Figure 7l
Material and locality. Single incomplete conch, SSMM
11601. From W8a, Wildenstein slice, Franconian Forest.
Tannenknock Formation, Wildenstein Member.
Description and discussion. A single incomplete internal
mould of a conch of ca. 2 mm height appears to represent
a species of Yochelcionella. This specimen resembles the
specimens of Leptostega irregularis in being slightly lat‑
erally compressed and by having two relatively prominent
plicae on the lateral sides. However, these ribs clearly fade
toward the anterior and posterior margins and do not form
recognizable ribs on these narrower fields of the conch. In
addition, the specimen lacks a clearly rearward directed
apex. Instead, the apex (although poorly preserved in the
specimen) appears to have been developed as a short,
slightly anteriorly directed conical structure. A relatively
broad tubular structure points into the posterodorsal direc‑
tion. This tube is apparently incompletely preserved, but
seems to have been short.
The exhalant tube is taken as a diagnostic character of the
genus Yochelcionella. The species possesses an impressive
variation of the morphology of the conch. Most of the 14
established species are characterized by a spirally enrolled
conch with the exhalant tube (“snorkel”) originating from
a part of the shell located well distant from the apical por‑
tion (see Atkins and Peel 2008 for an updated overview).
13

228
Fig. 11  Pelagiella sp. A. a, b, MMUW 2014A-140b, incomplete
conch, internal mould, Tannenknock Formation, Wildenstein Mem‑
ber, sample locality W18a, different view to the exterior of the ter‑
minal whorl. c, d, MMUW 2014A-140c, incomplete conch, inter‑
nal mould, Tannenknock Formation, Wildenstein Member, sample
locality W18a, views of the apical portion. e, MMUW 2014A-140d,
incomplete conch, internal mould, Tannenknock Formation, Wil‑
denstein Member, sample locality W18a. f, MMUW 2014A-175a,
incomplete conch, internal mould, Tannenknock Formation, Wilden‑
stein Member, sample locality W8. All scale bars 1 mm
The form from the Franconian Forest, however, is charac‑
terized by a tube branching off from near the apex with a
pseudo-exogastric coiling. The only formally described spe‑
cies of Yochelcionella with a pseudo-exogastric-type short
apical portion is Yochelcionella ostentata Runnegar and
Jell, 1976. This species from the late early Cambrian (Ord‑
ian) Coonigan Formation of New South Wales, Australia,
has numerous prominent and sharp plicae that completely
surround the conch with a broadly elliptical transverse sec‑
tion. A similar species is Yochelcionella gracilis Atkins and
Peel, 2004, with similar delicate, but prominent plicae and
a broadly subelliptical transverse section. This species from
latest early Cambrian lower part of the Henson Gletscher
Formation of northern Greenland has a somewhat extended
13
G. Geyer et al.
thin apical portion that varies between a pseudo-exogastric
and an endogastric coiling.
Another roughly similar species is Yochelcionella? erecta
(Walcott, 1891) from the Avalonian southeastern Newfound‑
land, which is clearly distinguished by numerous low con‑
centric rugae and an apex with a slightly endogastric direc‑
tion despite of the adjacent branching of the snorkel (see
Atkins and Peel 2008 for a detailed description).
A minute anteriorly directed apical portion is also known
from a single specimen from the Lemdad Syncline of the
High Atlas, Morocco, identified as “Genus novum et species
nova E” in Geyer (1986, pl. 4, fig. 61). The Moroccan speci‑
men, almost coeval with the specimen from the Franconian
Forest, is fairly similar, but differs in that it has numerous
low concentric ribs and a subcircular cross-section of the
conch. It should be emphasized that the late stratigraphic
occurrence of those pseudo-exogastric forms of Yochel-
cionella appears to confirm the assumption of Runnegar
and Jell (1976, p. 129) regarding an evolutionary trend with
a resulting morphocline from Latouchella-type species via
Y. penecyrano Runnegar and Jell, 1976 to pseudo-exogastric
species.
Order Pelagielliformes MacKinnon, 1985
[nom. trans. Parkhaev, 2001; ex Pelagiellida MacKinnon,
1985]
Diagnosis. Archaeobranchia with turbospiral shell; aperture
oval, elongated perpendicular to the coiling axis of the shell;
mantle caecum probably absent.
Family Pelagiellidae Knight, 1952
Genus Pelagiella Matthew, 1895
Type species. Cyrtolithes atlantoides Matthew, 1894 (by
original designation); Lower Cambrian of Hanford Brook,
New Brunswick, Canada.
Remarks. Innumberable studies on Cambrian small shelly
fossils have inflated the number of the species assigned tor
Pelagiella Matthew, 1895. At least 28 species have been
named (see a fairly comprehensive list in Parkhaev, 2001),
but in many cases the morphological plasticity and the diag‑
nostic characters are less than insufficiently known. Unfor‑
tunately, a necessary revision of these species would not
only require a reassessment of the original material, but in
some cases will need to be based on additional and better
preserved material.
Pelagiella sp. A
Figure 11
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
Material. Five specimens. From W18a: MMUW
2014A-140b, 2014A-140c, 2014A-140d; from W8: MMUW
2014A-175a.
Localities and stratum. Wildenstein slice, Franconian Forest,
sample localities W8 and W18a. Tannenknock Formation,
Wildenstein Member.
Description. Small, up to ca. 2.5 mm wide and 1.5 mm
high, dextrally coiled shell composed of 1.5‒2 moderately
expanding whorls; spire of adult individuals lies approxi‑
mately at the level of the upper apertural margin; initial
part of the juvenile whorl with elliptical transverse section,
barely expanding; last whorl wide and broad, with oval to
subtriangular transverse section near the aperture; suprape‑
ripheral part of the last whorl flattened; umbilicus narrow
and shallow; surface of internal mould without recognizable
microsculpture in the studied specimens; fine axial striation
corresponds to the growth lines of the shell.
Remarks. The preservation of the few available specimens
from the Wildenstein Formation are insufficient to character‑
ize this species. The largest specimen has a maximum diam‑
eter of the shell of just more than 2 mm, and the fracturable
preservation in strongly weathered sandy marlstone does not
allow a more complete preservation. It is noteworthy that
the two specimens that present the initial whorl (Fig. 11c,
d) show distinct differences in the width and arrangement
of the spire, but this may be attributable to the preservation
and morphologic plasticity.
The species is most reminiscent of Pelagiella subangu-
lata (Tate, 1892) but without a complete set of characters
that would allow a confident determination. Pelagiella atla-
sensis Geyer, 1986 from more or less coeval strata in the
Moroccan High Atlas shows similarly small conchs, but dif‑
fers in that it has gently convex initial whorls on the supra‑
peripheral side, and appears to have a more slender elongate
transverse section near the aperture.
Class Stenothecoida Yochelson, 1968
Order Cambridioida Horný, 1957
Family Cambridiidae Horný, 1957
Genus Stenothecoides Resser, 1938
Type species. Stenotheca elongata Walcott, 1884
Remarks. The enigmatic genus Stenothecoides is tradition‑
ally assigned to the Mollusca, but recent, largely unpub‑
lished studies emphasize similarities to the Brachiopoda.
Nevertheless, Stenothecoides is an unusual faunal element
with a surprisingly wide geographic range. It is primarily
distributed in Laurentia (e.g., Rasetti 1957; Savarese and
229
Signor 1989; Peel 1998), but also known from the Cambrian
continents Avalonia (e.g., Cobbold 1921, Landing 1991),
Kazakhstania (Koneva 1976, 1979a, b; Missarzhevsky and
Mambetov 1981; Popov and Holmer 1996), the Siberian
Foldbelts (Missarzhevsky and Mambetov 1981, Pel’man
et al. 1992), the North China Platform (Yu 1986, 1996) and
South China/Yangtze Platform (Qian et al. 1979; Yu 1996).
The specimen described below is the first report of the genus
from West Gondwana.
Stenothecoides? sp. indet.
Figure 12
Material. Single valve, internal mould, MMUW
2014A-156a.
Localities and stratum. Wildenstein slice, Franconian Forest,
locality W8. Tannenknock Formation, Wildenstein Member.
Description and discussion. The single present specimen is
a poorly preserved internal mould of a valve with a vague
indication of folds, and shows the slightly asymmetric out‑
line of the valve and direction characteristic for Stenothe-
coides. The apex lies marginal and is moderately acute and
not recognizably curved, forming the end of a moderately
prominent fold or keel which stretches nearly in the middle
of the valve, with a faint concavity to the dextral side, fading
towards the abapical end. The abapical margin is slightly
subtruncate obliquely.
The specimen is distinguished from most of the numerous
described species by its superficially symmetrical outline
and the obliquely subtruncate abapical margin in combi‑
nation to the subacute apical part. This feature is not the
product of imperfect preservation as indicated by the faintly
preserved growth lines that reflect ontogenetically earlier
shapes of the valve.
A superficial similarity exists with other genera of the
cambridiids, such as Bagenovia Horný, 1957 and Katun-
ioides Aksarina in Aksarina and Pel’man, 1978, but the rel‑
evant species of these genera have valves of diamond-like
shapes rather than being moderately elongate such as the
specimen from the Tannenknock Formation.
Phylum uncertain
Class Hyolitha Marek, 1963
Order Hyolithida Sysoev, 1957
Suborder Hyolithina Sysoev, 1957
[nom. transl. Geyer, 2018, ex Hyolithida]
Family Angusticornidae Sysoev, 1968
Remarks. Members of the family Angusticornidae are rec‑
ognized primarily by the morphology of the dorsum having
13

230
Fig. 12  Stenothecoides? sp. indet. a–d, MMUW 2014A-163a, incom‑
plete valve, internal mould, dorsal, anterior, oblique posterior and
lateral views. Tannenknock Formation, Wildenstein Member, sample
locality W8a. Scale bars 1 mm
a median carina, and by the planar to slightly convex venter
(Qian and Xiao 1995).
Genus Grantitheca Malinky, 1989
Type species. Grantitheca glenisteri Malinky, 1989; from the
lower Cambrian (Series 2, Dyeran) of the Taconic Alloch‑
thon of eastern New York State.
Diagnosis. Hyolithidae with angular longitudinal axis and
flat, steeply dipping adjacent slopes; lateral margins nar‑
rowly roundend and ligula short; aperture orthogonal with‑
out flare.
Grantitheca? klani sp. nov.
Figures 13, 14
Etymology. Named after the late Hermann Klan, a dedicated
collector of fossils from the Tannenknock Formation.
Holotype. MMUW 2014A-24, external mould of ventral side
and preserved part of internal mould of dorsum with septa.
13
G. Geyer et al.
Type locality. Locality W12 (on Fig. 2), classical Galgen‑
berg locality between Wildenstein and Premeusel villages,
Franconian Forest.
Type stratum. Galgenberg Member, Tannenknock Formation.
Studied material. Nearly two dozen partial conchs pre‑
served as external and internal moulds of the ventral sides,
partly with preserved parts of the dorsum, four opercula.
Paratypes. From sample horizon W8: MMUW 2014A-019;
from sample locality W12: MMUW 2014A-023, MMUW
2014A-065a, MMUW 2014A-065b, SSMM 10754, SSMM
10769; from sample locality W13: MMUW 2014A-062;
from W13a: MMUW 2014A-059, SSMM 10199, SSMM
10200, SSMM 10201, SSMM 10203, SSMM 10204, SSMM
10205, SSMM 10206, SSMM 10212; from sample horizon
W14a(?): SSMM 10171, SSMM 10172 (operculum), SSMM
10173 (operculum), SSMM 10186a (operculum), SSMM
10186b (part of internal mould of venter and external mould
of dorsum), SSMM 10789a (mould of dorsum with septa),
SSMM 10789b (mould of internal side of operculum);
from sample horizon T2: MMUW 2014A-043, MMUW
2014A-044.
Material tentatively assigned to Grantitheca? klani.
SSMM 10328, SSMM 10329, SSMM 11011 (from sam‑
ple horizon W8); SSMM 10198a, SSMM 10202, SSMM
10210, SSMM 10211, SSMM 10214 (all from sample hori‑
zon W13a); SSMM 10170 (from sample horizon W14a?).
Localities and stratum. Wildenstein slice, Franconian For‑
est. Tannenknock Formation, all from Galgenberg member
except for specimens from W8 (= Wildenstein Member).
Diagnosis. Species tentatively assigned to Grantitheca with
orthocone conch; flat venter and rounded dorsum; ventro-lat‑
eral muscle tracks developed along each edge of the venter;
cross-section elliptical to slightly subtriangular; venter cov‑
ered with transverse elements that correspond to the shape of
the ligula. Operculum monoclaviculate with pair of narrow
clavicles; cardinal processes short and almost perpendicular
to the clavicles.
Description. Hyolithid with an apical angle of ca. 35°. Maxi‑
mum size of complete specimens ca. 30 mm. Nearly flat
ventral side with fine growth-lines. Length of ligula ca. one
third of its width. Cross-section of the conch subtriangular
with rounded edges to almost elliptical. Dorsum rounded;
one lateral furrow along each edge of the dorsum is devel‑
oped. Apical region preserved; apical septa (usually seven
or eight) clearly visible and distinct (Figs. 13h, 14a). The
venter is covered with transverse rugae or lines that corre‑
spond with the shape of ligula. The dorsum bears indistinct
transverse lines.
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
Monoclaviculate operculum with pair of narrow clavi‑
cles. Clavicles subequal in width; the angle of divergence
ca. 140°. Cardinal processes not well preserved, but nar‑
row, short and almost perpendicular to the clavicles. The
surface of internal side of operculum is smooth without any
ornamentation (Fig. 13o, 14f). The outer side of operculum
is covered with growth lines (Fig. 13n).
Remarks. The ornamentation and morphology of venter of
Grantitheca? klani sp. nov. strongly resembles the speci‑
mens of Nevadotheca whitei (Resser, 1938) figured in Sun
et al. (2017, fig. 8). This resemblance clearly documents the
problem with hyolith systematics based solely on the conchs
(see Malinky 1988, p. 221; Sun et al. 2017, p. 90) which can
easily lead to an inexact or erroneous taxonomic placement.
The preserved operculum of Grantitheca? klani shows
a morphology resembling that of the genus Buchavalites
Marek, 1975 because of the monoclaviculate operculum
with long narrow clavicles and cardinal processes almost
perpendicular to the clavicles. However, the cardinal pro‑
cesses in Buchavalites are long and horizontally flattened.
Buchavalites differs in an ornamentation composed of nar‑
row longitudinal ribs on both sides and transverse densely
spaced growth-lines (see Marek 1975, p. 67) and lacking the
longitudinal furrows on the dorsum.
Grantitheca? klani differs from the type species of the
genus, Grantitheca glenisteri Malinky, 1989, in that it has
a less vaulted dorsum and less steeply sloping flanks of the
dorsum. The operculum of the type species is unknown so
the comparison of opercula is not possible.
Specimen MMUW 2014A-024 (Fig.  14a) shows an
attachment disk of an unknown organism. Some specimens
have an internal phosphatic plug that represents the sedi‑
mentary infilling of the conch, which itself is perforated by
burrows of the ichnogenus Arachnostega Bertling, 1992 as
visible on Fig. 13a, f (see Fatka et al. 2011 for additional
information on Arachnostega and hyolithids).
Many of the specimens placed under the Grantitheca?
klani are poorly preserved, deformed and lack some of the
critical characters. The specimens shown on Fig. 13 vary
considerably and may partly be mistaken as representing
different species, but transitional forms exist that straddle
from longitudinally deformed (e.g., Fig. 13b–d) to nearly
undeformed specimens (e.g., Fig. 14a, b). The placement
of such specimen within this taxon is thus tentatively. Nev‑
ertheless, these specimens show enough similarities to be
ranked in this taxon.
Family uncertain
Genus Hexitheca Sysoev, 1972, emend. Malinky and Berg-
Madsen, 1999
231
Type species. Hyolithus teretiusculus Linnarsson, 1877; from
the middle Cambrian of Sweden.
Hexitheca? sp.
Figures 15
Material. Six conchs. From sample locality W8: SSMM
10321; from sample locality W12: MMUW 2014A-029a,
b (internal and external mould of conch, ventral side); from
sample horizon W14: MMUW 2014A-004; from sample
horizon W14a(?): SSMM 10158, SSMM 10162, SSMM
10164.
Localities and strata. Wildenstein slice, Franconian Forest.
Galgenberg Member of Tannenknock Formation.
Description. Medium-sized conches with small apical angle
of ca. 14°–16°. The apical part is missing, but the original
size of the specimens can be reconstructed as more than
20 mm in length. Only the slightly vaulted ventral side is vis‑
ible indicating a comparatively short ligula. The ventral side
is partially cracked longitudinally and deformed owing to
compression during diagenesis. The sculpture of the conch
consists of regularly spaced transverse ribs (generally 6 to
8 per millimetre; 10 per millimetre in the apertural region).
Muscle scars are not visible.
Discussion. The ventral side of the conch resembles that
seen in Hexitheca teretiusculus (Linnarsson, 1877) in hav‑
ing distinct transverse sculpture; H. teretiuscula is a spe‑
cies known from the middle Cambrian (Eccaparadoxides
oelandicus Stage, E. insularis and Ptychagnostus praecur-
rens zones) of Sweden. Taxonomic placement within the
genus Hexitheca Sysoev, 1972 (emended Malinky and Berg-
Madsen 1999) is based herein only on the morphology of the
ventral side of conch. This venter is flat and not inflated as is
typical for Hexitheca, and the transverse sculpture of Hex-
itheca? sp. is more widely spaced and more prominent and
coarsely developed. The main similarities of the described
specimen with Hexitheca are the transverse sculpture, the
presumably short ligula and a similar apical angle of conch.
Other species that show similarities with Hexitheca? sp.
include Jincelites vogeli Valent et al., 2009 from the Dru‑
mian Jince Formation of Bohemia (see Valent et al. 2009,
figs. 3F–I) and Nevadotheca tenuistriata (Linnarsson, 1871)
that ranges from the late middle Cambrian Paradoxides
forchhammeri Stage (Solenopleura brachymetopa Zone) to
the Paibian Olenus Zone of Sweden (see Berg-Madsen and
Malinky 1999, fig. 5K).
Genus Jincelites Valent, Fatka, Micka and Szabad, 2009
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232 G. Geyer et al.

13
 Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
◂Fig. 13  a–k, m–o, l? Grantitheca? klani sp. nov. a SSMM 10754,
conch, ventral view of internal mould showing partially phosphatized
internal mould with corroded branched traces (Arachnostega); sam‑
ple locality W12; b SSMM 10199 conch, ventral view of exterior,
slightly dorsoventrally compressed; sample locality W13a; c SSMM
10212, conch, ventral view; sample locality W13a; d SSMM 10206,
ventral view of dorsoventrally flattened conch with imperfectly pre‑
served axial vessel (brown); sample locality W13a; e SSMM 10171,
ventral view of dorsoventrally flattened conch with impression of
apertural margin of dorsal side; sample locality W14a?; f SSMM
10739, incomplete conch with partially phosphatized steinkern pen‑
etrated by branched traces (Arachnostega); sample locality W12; g
SSMM 10204, conch, ventral view of exterior, dorsoventrally com‑
pressed; sample locality W13a; h SSMM 10769, conch, ventral view
of infilling with seven preserved septa in apical part and stratification
in the remaining chamber; sample locality W12; i MMUW 2014A-
001, ventral view of dorsoventrally flattened conch; sample locality
W11; j SSMM 10203, ventral view of dorsoventrally flattened conch;
sample locality W13a; k SSMM 11011, partial conch, apical view of
phosphatized filling showing transverse profile; sample locality W8; l
SSMM 10251, phosphatized partial internal mould of conch with pre‑
served traces; sample locality W13a; m SSMM 10198a, conch, dorsal
view of semi-three-dimensionally preserved internal mould; sample
locality W13a; n SSMM 10173, incomplete operculum, apertural
view; sample locality W14a?; o SSMM 10172, operculum, view of
internal face; sample locality W14? All specimens from the Galgen‑
berg Member of the Tannenknock Formation. Scale bars equal 5 mm,
except in i and k (= 1 mm)
Type species. Jincelites vogeli Valent et al., 2009; from
the middle Cambrian of the Příbram-Jince Basin, Czech
Republic.
Jincelites cf. vogeli Valent, Fatka, Micka and Szabad, 2009
Figure 16
Studied material. Six conchs. From sample horizon W9:
SSMM 10346; from W12: SSMM 10767; from sample hori‑
zon W14a?: SSMM 10187a, b, SSMM 10188; from sample
horizon T2a: SSMM 10038.
Localities and strata. Wildenstein and Triebenreuth slices,
Franconian Forest. Galgenberg Member of Tannenknock
Formation except for W9 (Wildenstein Member).
Description. Hyolithid with orthocone conch, apical angle
ca. 12°; cross-section of conch probably subtriangular. The
ligula is short and moderately rounded; the length of ligula
attains ca. 40% of the width of the conch. An axial keel is
developed on the dorsum. The ornamentation of the venter
and dorsum consists of the distinct and regularly spaced ribs.
Discussion. The specimens closely resemble Jincelites
vogeli Valent et al., 2009 from the Drumian Jince Forma‑
tion of Bohemia (see Valent et al. 2009, fig. 3F–I) by the
morphology and ornamentation on the venter and dorsum
and by a similarly shaped ligula. They differ in the apical
angle of conch (12° in Jincelites cf. vogeli; 20° in J. vogeli).
233
A confident determination of the present specimens is not
possible without knowledge of the operculum.
Genus Maxilites Marek, 1972
Type species. Maxilites snajdri Marek, 1972; from the mid‑
dle Cambrian of the Příbram–Jince Basin, Czech Republic.
Maxilites? sp.
Figure 17
Material. One specimen, SSMM 10391, external mould of
conch. From Wildenstein slice, Franconian Forest, locality
W9. Tentatively assigned to the same form: SSMM 10326,
SSMM 10326, from sample horizon W8. All from Tannen‑
knock Formation, Wildenstein Member.
Description. Orthocone conch with highly inflated dorsum;
venter unknown. The surface of the conch covered with
very fine and slightly indistinct lines. Apical angle ca. 32°.
Length of the conch ca. 30 mm.
Discussion. The hyolithid represented by the conch is tenta‑
tively placed under the genus Maxilites Marek, 1972 based
on the large, rapidly expanding and robust conch and the
highly inflated dorsum, which is covered with fine lines. A
more precise comparison with the species of Maxilites is
impossible until the becomes known.
Hyolithid genus and species A
Figure 18b
Material. Single operculum, external mould of internal side,
MMUW 2014A-081.
Locality and stratum. Wildenstein slice, Franconian Forest,
locality W8. Tannenknock Formation, Wildenstein Member.
Description. The operculum is biclaviculate, with the ante‑
rior margin of the inner side preserved. The area of the oper‑
culum where the cardinal processes are located (cardinal
margin) is damaged and partly missing. The angle of diver‑
gence of the first pair of very narrow clavicles attains 130°;
the second pair of clavicles includes an angle of divergence
of ca. 115°. The overall shape of the operculum cannot be
determined because the cardinal margin is not present. No
sculpture or muscle scars are visible.
Discussion. The single preserved operculum is the first bicla‑
viculate operculum known from Cambrian strata. It resem‑
bles Decipilites decipiens (Barrande, 1867) from the Katian
(late Ordovician) of Bohemia. This species of the family
Pauxillitidae Marek, 1967 has a biclaviculate operculum
13

234 G. Geyer et al.

13
 Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
◂Fig. 14  a–f, h–n, g? Grantitheca? klani sp. nov. a MMUW 2014A-
024, conch, ventral view of partial internal mould and exfoliated
dorsal part with apical septa, with attachment disc of unknown
organism; sample locality W12; b MMUW 2014A-066, conch, ven‑
tral view of internal mould and exfoliated with apical septa; sample
locality W12; c MMUW 2014A-059, conch, dorsal view of partial
internal mould and exfoliated ventral part; sample locality W13a; d
SSMM 10205, conch, ventral view with partly phosphatized internal
mould and numerous growth lines; sample locality W13a; e SSMM
10186a, ventral view of crushed conch with distinct growth lines;
sample locality W14a?; f SSMM 10789a, operculum, apertural view
of internal mould; sample locality W14a?; g MMUW 2014A-061,
partial conch with partial phosphatized internal mould with Arach-
nostega-type traces and compacted shell with growth lines (right);
sample locality W8; h, i SSMM 10186b, conch, ventral view with
partial phosphatized internal mould (h) and magnification showing
transverse rugae (i); sample locality W14a; j MMUW 2014A-019,
fragment of conch, ventral view of internal mould; sample local‑
ity W8; k MMUW 2014A-065a, incomplete conch, ventral view of
internal mould; sample locality W12; l MMUW 2014A-065b, incom‑
plete conch, dorsal view of external mould; sample locality W12;
m MMUW 2014A-044, incomplete conch, dorsal view of exter‑
nal mould; sample locality T2;. n MMUW 2014A-043, incomplete
conch, ventral view of internal mould; sample locality T2. All speci‑
mens from the Galgenberg Member of the Tannenknock Formation.
Scale bars equal 5 mm, except in f and g (= 1 mm)
with narrow clavicles, but differs distinctly in that it has a
much more obtuse angle of divergence of the clavicles and
in the overall shape of the operculum in which the poste‑
rior part of operculum suggests shorter ligula. Biclaviculate
opercula are rare and have been reported only from the late
Ordovician to early Devonian (Marek 1967, p. 61).
Hyolithid genus and species B
Figure 18a, c–e
Material. Three specimens, SSMM 10764 (deformed inter‑
nal mould of venter and part of external mould of dorsum),
SSMM 10765 (external mould of venter with septate apical
part) and SSMM 10766. Tentatively assigned to the same
form: SSMM 10773 (slightly crushed conch).
Localities and stratum. All specimens from W12, classical
locality of Galgenberg Member, Tannenknock Formation,
Galgenberg, Wildenstein slice, Franconian Forest.
Description. Hyolithid with sharp apical angle of 15°–18°.
The ligula is short, its length ca. half width of the aper‑
ture. With 7–8 septa in the apical part of the conch. Visible
dorsum in SSMM 10764 (Fig. 19a) broken along its longi‑
tudinal axis. Dorsum probably subtriangular in transverse
profile.
Remarks. Important characters such as the morphology of
the operculum and the sculpture are missing so that an accu‑
rate taxonomic determination is not possible. The specimens
235
are classified into the order Hyolithida because of the pres‑
ence of a ligula.
Suborder Aladracina Geyer, 2018
Discussion. The genus Aladraco Geyer, 2018, formerly
known as Oxyprymna Kiderlen, 1933, a preoccupied generic
name, bears morphological characters such as the size, shape
and external ornamentation of the conch, which indicate a
systematic position in the class Hyolitha. A number of char‑
acters, such as a tripartition into an axial chamber and lateral
acuminate processes, and the course of the apertural margin,
differ considerably from those seen in the two orders Hyo‑
lithida and Orthothecida, and suggest distinct differences
in functional morphology. However, similarities of the
morphological characters with the known representatives
of the Hyolithida advocate that the genus Aladraco and its
(mostly undescribed) relatives are derived from this system‑
atic group and should be dealt with as a hitherto unrecog‑
nized distinct suborder and family, which was introduced as
Aladracina and Aladracidae by Geyer (2018). An additional
genus of the suborder is described below under the name
Cambrachelous gen. nov.
Family Aladracidae Geyer, 2018
Genus Aladraco Geyer, 2018
(pro Oxyprymna Kiderlen, 1933, p. 166, non Oxyprymna
Stål, 1893, p. 5).
Type species. Conularia schloppensis Wurm, 1925 (by origi‑
nal monotypy; Kiderlen 1933).
Remarks. The name Aladraco has been suggested recently
(Geyer 2018) to replace the preoccupied name Oxyprymna
used by Kiderlen (1933) when he erected the genus. Two
species, Aladraco schloppensis (Wurm, 1925) and A. oug-
natensis sp. nov. are recognised from coeval strata of the
traditional lower (or lowermost) middle Cambrian of West
Gondwana. For differences to Cambrachelous gen. nov. (see
below).
Aladraco schloppensis (Wurm, 1925b)
Figure 19
v * 1925 Conularia Schloppensis nov. sp.—Wurm, p. 77,
pl. 3, figs. 1, 2.
v 1925 Conularia sp. b—Wurm, p. 77, pl. III, fig. 3.
1933  Oxyprymna schloppensis Wurm sp.—Kiderlen,
pp. 166–172, figs. 1–5, 7, ?14.
v 2018 Aladraco schloppensis (Wurm, 1925)—Geyer,
pp. 86, 87, 88, 90, 93, 94, 97–98, figs. 2–4, 6.
13

236
Fig. 15  Hexitheca? sp. a MMUW 2014A-029a, incomplete conch,
partly exfoliated; sample locality W12; b, f MMUW 2014A-029b,
incomplete conch, partly exfoliated, detail in F shows expression of
concentric rugae in flattened conch; sample locality W12; c, g SSMM
10321, incomplete conch with partly phosphatized shell; weakly min‑
eralized apertural end in g shows expression of concentric rugae in
flattened conch; sample locality W8; d SSMM 10162, incomplete
conch with partial internal mould preserved as phosphatized infilling
and external mould of apical half with weakly preserved rugae; note
See Geyer (2018) for additional synonymy.
Material. Ca. 35 conchs (see Geyer 2018 for list).
13
G. Geyer et al.
staining of rock enveloping the middle part of the conch and possibly
resulting from dispersal of organic matter; sample locality W14a?; e
SSMM 10164, incomplete conch with partial phosphatized internal
mould penetrated by traces; sample locality W14a?; h, SSMM 10158,
incomplete conch with crushed, mineralized shell; sample locality
W14a?; i MMUW 2014A-004, partial conch with crushed shell; sam‑
ple locality W12. All specimens from the Galgenberg Member of the
Tannenknock Formation. Scale bars equal 5 mm in a–e, h, 1 mm in f,
g and i 
Localities and strata. All specimens from the Wildenstein
slice, localities W8, W9, W16b and W17 (Wildenstein
Member), W12, W12c and W13a (Galgenberg Member),
Tannenknock Formation, Agdzian Stage.
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
Fig. 16  Jincelites cf. vogeli Valent, Fatka, Micka and Szabad, 2009.
a SSMM 10188, conch, ventral view with axially crushed shell; note
widely spaced growth lines near apertural margin; from W14a?; b
SSMM 10187a, b, two incomplete conchs, partly crushed, ventral
views; from sample horizon W14a?; c SSMM 10346, conch, ventral
view showing growth rhythms; from sample horizon W9; d SSMM
Discussion. Aladraco schloppensis (Wurm, 1925) is the only
named hyolith species described in the original description
of the Tannenknock fauna in Wurm (1925). It is also the
largest and most conspicuous species, known from speci‑
mens of up to 65  mm in length. A detailed description
and discussion of the species, as well as its synonymy was
provided by Geyer (2017b) so the species is only included
here for completeness with photographs to illustrate its
morphology.
237
10767, partial conch with partly phosphatized internal mould; from
sample locality W12; e SSMM 10038, conch, internal mould with
delicate, densely spaced growth lines; from sample horizon T2a. All
specimens from the Galgenberg Member of the Tannenknock Forma‑
tion except for C (from Wildenstein Member). Scale bars equal 5 mm
in A–D, 1 mm in E
Cambrachelous gen. nov.
Type species. Cambrachelous diploprosopus gen. et sp. nov.,
from the Tannenknock Formation, Stage 4–Wuliuan bound‑
ary interval, Franconian Forest, Germany.
Etymology. Named after its occurrence in the Cambrian and
after the Greek mythologic figure Acheloos (Ἀχελώїoς in
ancient Greek), chief of all river deities, capable of changing
13

238
Fig. 17  Maxilites? sp. SSMM 10391, incomplete conch, oblique dor‑
sal views. Wildenstein Member of the Tannenknock Formation, sam‑
ple locality W9. Scale bar 5 mm
his shape; an allusion to the change of the dorsal profile of
the conchs from small to large specimens.
Diagnosis. Genus of the Aladracidae characterised by a
moderately large conch with an apertural angle of ca. 20°–
25°; dorsum divided into a nearly flat median sector defined
by two moderately prominent keels and a slightly sunken
area between them, lateral sectors sloping abaxially; a third
keel in axial position developed only in juvenile and up to
middle-sized individuals. Apertural margin planar.
Discussion. The new genus Cambrachelous is typified by
a tripartite subdivision of the dorsal and ventral sides of
the conch, with a distinctly raised median/axial sector on
the ventral side so that the lateral parts of the conch form
relatively acuminate extensions as seen in Aladraco Geyer,
2017. Together with the planar aperture this characterises
Cambrachelous as a second genus of the recently introduced
suborder Aladracina.
The differences from Aladraco are considerable when
in terms of functional aspects. Aladraco has a more
clearly raised central sector on the dorsal side and a cen‑
tral sector with a ventrally raised surface on the venter
which result in a fairly well defined central chamber. The
central sector on the dorsum of Cambrachelous is moder‑
ately raised only, and in the gerontic stages of the conches
this sector is more or less clearly sunken medially. On the
dorsum, the central sector is more or less flat. As a result,
Cambrachelous does not have a well-defined functional
13
G. Geyer et al.
axial chamber, but nonetheless shares the relatively nar‑
row lateral extensions seen in Aladraco. A comparison of
the transverse sections of both genera is shown in Fig. 21.
Cambrachelous is further characterized by a smaller api‑
cal angle of ca. 20°–25° and by the development of keels
on the dorsum.
Cambrachelous superficially resembles the monotypic
orthothecid genus Probactrotheca Marek in Valent et al.,
2012, known only from P. briketa Valent et al., 2012 from
the Buchava Formation, Eccaparadoxides pusillus Biozone,
Drumian, of Bohemia. Probactrotheca briketa is character‑
ized by a pronounced axial keel on the dorsum flanked by
sunken areas and with slightly inflated dorsolateral portions
that tend to be subangular. Its ventral side is concave, with
a broad median depression and raised shoulders (when seen
from ventral aspect). This provides a quite distinctive cross
section (Valent et al. 2012, fig. 3) that is derived from the
trapezoidal shape, and the relatively thin lateral portions
seen in Cambrachelous are lacking. The apertural margin
of P. briketa appears to have been more or less planar as in
Cambrachelous.
Cambrachelous diploprosopus gen. et sp. nov.
Figure 20, 21
Etymology. From the Greek διπλός, double, and πρόσωπο,
face, referring to the change in the dorsal appearance of the
conch by reduction of keels.
Holotype. SSMM 10567, fairly complete conch (Fig. 20i, j).
Type locality and type stratum. Wildenstein slice, Franco‑
nian Forest, Germany, sample locality W8. Tannenknock
Formation, Wildenstein Member, Cambrian Stage 4–Wuli‑
uan boundary interval.
Studied material. Ca. 20 conchs. Paratypes: From sam‑
ple horizon W8: MMUW 2014A-076, SSMM 10318,
SSMM 10323, SSMM 10567, SSMM 10568, SSMM
11142, SSMM 11143a, SSMM 11143b (counterpart of
11143a); from sample horizon W9: MMUW 2017B-016;
from sample horizon W11: MMUW 2014A-002, MMUW
2014A-025a, SSMM 10771; from sample horizon W11/
W12: MMUW 2014A-095; from sample horizon W12:
MMUW 2014A-099; from sample horizon W13a: SSMM
10252, SSMM 10253, SSMM 10254; from sample hori‑
zon W16b: MMUW 2014A-013a; from sample horizon T2:
SSMM 10280. Tentatively assigned to the same form: from
sample horizon W8: SSMM 10566, 11044; from sample
horizon W9: SSMM 10316.
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany 239

Fig. 18  a, c–e Hyolithid genus and species B. a SSMM 10764, ven‑ ▸

tral view of deformed conch; c SSMM 10765, fragment of conch,
apical septate part; d, e SSMM 10766, nearly complete conch, partly
exfoliated with septate apical portion and remains of phosphatized
infilling. All specimens from Galgenberg Member, Tannenknock
Formation, sample horizon W12. Scale bars equal 1  mm. b Hyo‑
lithid genus and species A, MMUW 2014A-081, operculum, internal
mould; Wildenstein Member, Tannenknock Formation, sample hori‑
zon W8. Scale bar 1 mm

Localities and strata. Wildenstein slice, Franconian For‑

est. Tannenknock Formation, Galgenberg and Wildenstein
Members.

Diagnosis. Diagnosis of genus (because of monotypy).

Description. Conch with apertural angle varying between

19° and 26° in the present specimens, with broader angles
caused by slight dorsoventral compression. Dorsal side
divided by two moderately prominent, slightly rounded keels
into a central and two lateral sectors. The middle sector is
narrow and flat or sunken between the principal keels. A
third keel in axial position is developed in the central sector
in juvenile and middle-sized conchs. The lateral sectors form
flanks that slope distinctly abaxial of the keels, but are more
or less flat for most of its stretch. The dorsal surface of the
conch is mostly smooth, but low and indistinct, rhythmic
transverse ribs on the lateral areas are visible in two of the
specimens, and one of them shows the ribbing on the keels
as well (Fig. 20g). A single specimen preserves remnants
of the ribs in the central area (Fig. 20h). The shape of these
ribs has probably been reinforced by taphonomic conditions.
The ventral side of the conch is difficult to precisely
characterize. However, careful investigation of the avail‑
able material suggests that the venter also shows a triparti‑
tion into a central and two lateral areas, which are separated
by relatively narrow bands oblique to the princinpal plane
that is defined by the lateral edges. Keels are absent when
exfoliated specimens or external moulds of the venter are
preserved. Such specimens are shown in Fig. 20e, f. The
specimen in Fig. 20a appears to be a composite mould pre‑
served in shale in which the dorsal is pressed on the ventral
mould of the same conch.

Discussion. Although no specimen is known that allows

in situ recognition of the cross-section, the combination of
dorsal and ventral surface indicates a situation comparable to
that known from Aladraco (see above): The species had an

13

240 G. Geyer et al.

13
 Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
◂Fig. 19  Aladraco schloppensis (Wurm, 1925a, b). a, b lectotype,
SNSB-BSPG-1924-XII-5, large conch, original of Conularia schlop-
pensis in Wurm (1925a, b, pl. III, fig. 5), almost certainly from local‑
ity W9; a dorsal view of entire conch, b detail of dorsum near aper‑
tural end showing faint transverse growth lines and narrow groove
on internal mould indicting thickened shell along longitudinal keel
(arrows); c MMUW 2017B-002, dorsum of large conch with partly
preserved longitudinal threads (arrow), dorsal view; from local‑
ity W12c; d MMUW 2017B-013, small specimen, oblique apertural
view of dorsal part of the shell with infilling of axial chamber par‑
tially preserved near apertural margin; from locality W9; e, f SNSB-
BSPG -1924-XII-502, large conch, mould of venter with remnants of
shell, dorsal (f) and oblique anterior views (e), almost certainly from
locality W9; original of Conularia schloppensis in Wurm (1925a, b,
pl. III, fig.  4); g MMUW 2017D-009, large conch, internal mould,
dorsal view; from locality W9; h MMUW 2017D-010; large conch,
ventral view with convex axial chamber; from locality W9. Scale bars
equal 5 mm in a, c–h (equivalent to magnification ×2 except for c),
1 mm in b 
axial compartment in which most of the cavity of the conch
was concentrated. The lateral “chambers” were conical in
transverse profile and slightly sloping ventrally so that the
base of the axial portion was above the lateral edges of the
conch (Fig. 21b). However, other than in Aladraco the cen‑
tral sectors did not form a distinct chamber with a ventrally
vaulted bottom, and the dorsal convexity in the central sec‑
tor was not as distinct in Cambrachelous diploprosopus as
in the species of Aladraco. Among the several other other
characters that distinguish Cambrachelous diploprosopus
from the species of Aladraco, the smaller apertural angle
and the absence of distinct longitudinal lirae are particularly
distinctive.
Order Orthothecida Marek, 1966
Family uncertain
Genus Brevitheca Marek, 1967
Type species. Brevitheca minimax Marek, 1967; from the
Caradocian, Late Ordovician, of Bohemia.
Brevitheca? sp.
Figure 22
Material. Two partial conches; MMUW 2014A-009,
MMUW 2014A-010a, b.
241
Locality and stratum. Wildenstein slice, Franconian Forest,
sample horizon W15a; Tannenknock Formation, Wilden‑
stein Member.
Description. Moderately large conch with preserved aper‑
tural part (apical part missing). Original total length of the
conch ca. 23 mm, apical angle probably ca. 22°. Ventral side
almost flat. Sculpture of ventral side consists of well defined,
closely spaced rounded ribs. Muscle scars not recognizable.
Discussion. Specimen MMUW 2014A-10 shows only the
ventral side with a distinct transverse sculpture. This sculp‑
ture resembles that seen in the genus Brevitheca Marek,
1967 from the early Katian Zahořany Formation (Upper
Ordovician) of Bohemia.
A precise taxonomic placement is not possible due to the
absence of data on the dorsum and the operculum, although
the features that are preserved suggest affinity to Brev-
itheca. Nevertheless, the specimen appears to represents a
yet unknown genus and species for the Cambrian of West
Gondwana.
Genus Nephrotheca Marek, 1966
Type species. Orthotheca sarkaensis Novák, 1891, from the
Šárka Formation, Darriwilian, Middle Ordovician, of the
Prague Basin, Czech Republic.
Nephrotheca? sp.
Figure 23
Material. Internal and external moulds of a single conch
with exposed ventral side; MMUW 2014A-067a, b.
Locality and stratum. Wildenstein slice, Franconian Forest;
sample horizon W12c, Galgenberg Member of Tannenknock
Formation.
Description. Conch with restored original length of almost
30 mm. Apical angle 12°. Ventral side almost flat, with only
faintly concave zone along the mid-axis. Sculpture of ventral
side consists of only indistinct longitudinal lines; longitu‑
dinal ornamentation on lateral side clearly visible. Cross-
section of the conch subtriangular with strongly rounded
13

242 G. Geyer et al.

13
 Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany 243

◂Fig. 20  Cambrachelous diploprosopus gen. et sp. nov. a SSMM corners. Apical part present, but no septa visible. Apertural
10318, paratype, conch, dorsal view of internal mould; sample hori‑ part missing, muscle scars not recognizable.
zon W8; b SSMM 10280, paratype, conch, dorsal view of internal
mould; from sample horizon T2; c MMUW 2017B-016, paratype,
conch, dorsal view of internal mould; from sample horizon W9; d Discussion. The cross-section and longitudinal ornamen‑
SSMM 10771, paratype, conch, dorsal view of internal mould with tation of the conch matches those known from the genus
triplicate dorsum; from sample horizon W12; e SSMM 10254, para‑ Nephrotheca Marek, 1966 (see recent remarks in Valent
type, conch, dorsal view of venter, external mould; from sample hori‑
zon W13a; f SSMM 10253, paratype, conch, dorsal view of internal
et al. 2013, p. 119–121). The specimens are also similar to
mould; from sample horizon W13a; g MMUW 2014A-095, paratype, species of the genus Decoritheca Sysoev, 1972. It is impos‑
slightly deformed conch, dorsal view of partly exfoliated specimen sible, however, to decide on the precise generic affinity
with clearly visible riblets on right-hand flank; from sample horizon because species of both genera may have a shallow ventral
W11/W12; h SSMM 10323, paratype, slightly deformed conch, dor‑
sal view of internal mould with faint riblets; sample horizon W8; i,
furrow, longitudinal ornamentation and a small apical angle
j SSMM 10567, holotype, conch, dorsal views of internal mould in (13° in Nephrotheca sophia Valent et al. 2013).The absence
coated (i) and uncoated (j) conditions; note development from tri‑ of the apertural part, however, prevents a confident place‑
carinate to bicarinate profile of dorsum; from sample horizon W8; k ment under this genus because it remains unclear whether
MMUW 2014A-099, paratype, conch of immature individual, dorsal
view; sample horizon W12; l MMUW 2014A-002, paratype, conch of
this specimen belongs to order Hyolithida or Orthothecida.
immature individual, dorsal view; sample horizon W11; m MMUW
2014A-025a, paratype, cracked conch of immature individual, dorsal Order uncertain
view; sample horizon W12; n MMUW 2014A-076, paratype, partial
conch, dorsal view of internal mould; sample horizon W8; o MMUW
Hyolith genus and species indeterminate A
2014A-013a, paratype, conch, latex cast of internal mould, dorsal
view of latex cast, detail showing polyplicate dorsum; sample horizon Figure 24a
W16b. Galgenberg and Wildenstein Members, Tannenknock Forma‑
tion. Scale bars equal 5 mm in a–j, n, o, 1 mm in k–m  Material. Single incomplete conch, external mould of ven‑
tral side, MMUW 2014A-013.

Locality and stratum. Wildenstein slice, Franconian Forest,

sample horizon W2. Tannenknock Formation, Galgenberg
Member.

Description. Hyolith of moderate size, estimated length ca.

20–25 mm. Characteristic sculpture on ventral side of the
conch consists of fine longitudinal lines which are more dis‑
tinct close to the lateral edges and almost invisible along the
central sector of the conch. Apical angle ca. 25°. Apertural
and apical part not preserved. Muscle scars not recognizable.

Discussion. The longitudinal ornamentation is the only rec‑

ognizable taxonomic character of this specimen, but this
type of ornamentation is common within the Hyolitha (e.g.,
Berg-Madsen and Malinky 1999; Malinky 2002).

Hyolith genus and species indeterminate B

Figure 24b–d

Fig. 21  Cross-sections of the conches of Aladraco schloppensis Material. Ca. 8 conchs, in repository: MMUW 2014A-28a, b
(Wurm, 1925a, b) (a) Cambrachelous diploprosopus gen. et sp. nov. (internal and external mould of single conch), SSMM 10740
(b), and Probactrotheca briketa Valent et al., 2012 (c) and SSMM 10741 (crushed partial conches).

13

244
Fig. 22  Brevitheca? sp. a MMUW 2014A-009, crushed fragment of conch. b, c MMUW 2014A-010a, b, fragment of conch; b partly exfoliated
ventral side; c counterpart of b. Both specimens from sample horizon W15a, Wildenstein Member, Tannenknock Formation. Scale bars 1 mm
Locality and stratum. Wildenstein slice, Franconian Forest,
sample horizon W12. Tannenknock Formation, Galgenberg
Member.
Description. Small-sized hyolith, orthoconic conch with api‑
cal angle of ca. 20°. Apical part subelliptical or subcircular
in transverse section. Apertural part flabellate, undulate on
venter, but mostly broken off. Muscle scars and sculpture
not recognizable.
Discussion. The more precise taxonomic placement is not
possible because the apertural part is missing. Thus, it is
impossible to ascertain whether a ligula was developed.
Hyolith genus and species indeterminate C
Figure 24e
Material. Single specimen, MMUW 2014A-058.
13
G. Geyer et al.
Locality and stratum. Wildenstein slice, Franconian Forest,
sample horizon W13a. Tannenknock Formation, Galgenberg
Member.
Description. Small-sized hyolith, length ca. 10 mm. Angle
of divergence ca. 20°. Apertural section missing. Dorsal
side highly convex, ventral side apparently slightly concave.
Muscle scars and sculpture not recognizable.
Discussion. This form is similar to Orthothecid genus and
species indeterminate A, but the missing apertural end pre‑
vents a taxonomic placement within the order Orthothecida.
Hyolith genus and species indeterminate D
Figure 25a, c
Material. MMUW 2014A-097, partial internal mould of
conch; MMUW 2014A-098, external mould of ventral side.
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany
Fig. 23  Nephrotheca? sp. MMUW 2014A-067a, external mould of
conch with ventrolateral side exposed; a, c latex cast of mould, entire
specimen and enlargement of area near apertural end; b external
mould with remains of shell. From sample horizon W12c, Galgen‑
berg Member, Tannenknock Formation. Scale bars equal 5  mm in a
and b, 1 mm in c 
Locality and stratum. Wildenstein slice, Franconian Forest,
sample horizon W11/12. Tannenknock Formation, Galgen‑
berg Member.
245
Description. Orthothecid conch with a length of 13 mm.
Apical angle 19°. The cross section of the conch is suboval
with ventral side less convex than the dorsal side; lateral
edges strongly rounded. The apical part is missing in the
available specimens. Septa, muscle scars and sculpture not
recognized.
Discussion. The circular to oval cross-section is very com‑
mon within the order Orthothecida. Many specimens or spe‑
cies with this shape were classified in the genus Circotheca
Sysoev, 1958 (emend. Berg-Madsen and Malinky, 1999).
The presence of this sole morphological character does not
allow any more precise taxonomic placement (see remarks
on Circotheca in Berg-Madsen and Malinky 1999, p. 873).
Hyolith genus and species indeterminate E
Figure 25b, d, e
Material. 2–4 conches. From sample horizon W12: MMUW
2014A-40a, b, internal and external moulds of dorsal side;
SSMM 10738, conch with phosphatic infilling. Tentatively
belonging to the same form, from sample horizon W12:
SSMM 10737; from sample horizon W14a?: SSMM 10159.
Locality and stratum. Wildenstein slice, Franconian Forest.
Galgenberg Member of Tannenknock Formation.
Description. Conch of orthothecid hyolith with a length of
l8 mm. Apical angle 18°. Cross-section of conch probably
subtriangular with sharply rounded lateral edges. Ventral
side not visible in the available specimens. Apical part miss‑
ing. Septa, muscle scars and sculpture not visible.
Discussion. The subtriangular cross-section of this specimen
is typical for the family Orthothecidae Sysoev, 1958 emend.
Malinky (2009, p. 591), and this feature clearly distinguishes
this form from Orthothecid genus and species indeterminate
B described before. Other important characters such as oper‑
culum and sculpture are missing preventing a more accurate
taxonomic placement.
13

246
Fig. 24  a Hyolith genus and species indeterminate A, MMUW
21014A-013, incomplete conch, external mould of ventral side;
sample horizon W2. Scale bar 5  mm. b–d Hyolith genus and spe‑
cies indeterminate B. b SSMM 10743, crushed partial conch; sam‑
ple horizon W12;. c MMUW 21014A-028b, incomplete conch, partly
Fig. 25  a, c Hyolith genus and species indeterminate D. a MMUW
2014A-098, conch, external mould; sample horizon W11/12; c
MMUW 2014A-097, partial internal mould of conch, cross-section;
sample horizon W11/12. Scale bars 1  mm. b, d, e Hyolith genus
and species indeterminate E. b SSMM 10738, conch, phosphatized
13
G. Geyer et al.
exfoliated, dorsal view showing undulations towards aperture; sample
horizon W12; d MMUW 21014A-028a, incomplete crushed conch,
counterpart of MMUW 2014A-028b. e Hyolith genus and species
indeterminate C, MMUW 2014A-058, incomplete conch, internal
mould of dorsal side; sample horizon W13a. Scale bars 1 mm
internal mould with trace fossils; sample horizon W12; d MMUW
2014A-40b, conch, internal mould; sample horizon W12;. e, MMUW
2014A-40a, conch, external mould as counterpart of MMUW 2014A-
40b in d; sample horizon W12. Scale bars 1 mm
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany 247

Acknowledgements  This study includes almost all available mollusc
specimens known from the Tannenknock Formation. The material
comes partly from the collection of Stefan Meier (Marktredwitz), other
specimens come from collections of the late Klaus Sdzuy (Würzburg)
and the senior author. Loan of specimens was made possible by Martin
Nose and Winfried Werner (Bayerische Staatssammlung für Geologie
und Paläontologie, Munich) and Erwin Geiß (Bayerisches Landesamt
für Umwelt, Augsburg/Munich). We thank Anna Żylińska and Bożena
Plejznerowska (both Warsaw University) for help accessing literature.
The material used in this study was partly collected in the course of
several research grants of the Deutsche Forschungsgemeinschaft (DFG)
to the senior author, and the preparation of the manuscript was made
possible by research grant GE 549/22-1 to GG. This work was finan‑
cially supported by the Ministry of Culture of the Czech Republic
(DKRVO 2018/06, National Museum, 00023272) to MV. We gratefully
acknowledge comprehensive reviews by John Malinky (Escondido, CA,
USA), Sarah Jacquet (University of Missouri, Columbia, MO) and
Pavel Yu. Parkhaev (Russian Academy of Sciences, Moscow, Russia),
which significantly helped to improve this article.
localities indicated in Fig. 2. Occurrences of Helcionelloida,
Pelagiellida, Stenothecoida, and Hyolitha according to the
presented data, trilobite data from Geyer (2017).
Collectors are (in alphabetical order): Erwin Albert,
Burgkunstadt (†); Josef Gandl, Würzburg; Gerd Geyer
(author); Wolfgang Hammann, Würzburg (†); Peter Hick‑
ethier, Zedtwitz; Jörg Hiltl, Erlangen; Reverent S. G. Kohl‑
mann, Stadtsteinach (†); Hermann Klan, Hof (†); Ed Land‑
ing, Albany, NY, USA; Volkmar Ludwig, Würzburg (†);
Stefan Meier, Marktredwitz; Bruno Paulus, München (†);
Rudolf Richter, Frankfurt a. M. (†); Armin Rückert, Würz­
burg/Neuried; Klaus Sdzuy, Würzburg (†); Harald Trage‑
lehn, Köln/Wallenfels; Wolfgang Trapp, Würzburg; and
Adolf Wurm, Würzburg (†). Some specimen in the SMF
repository were collected by unidentified members of the
University of Frankfurt a. M.

Appendix: Samples from Tannenknock

formation relevant for this study

Locality IDs starting with “W” or “T” and with Arabic

numbers refer to occurrences in the Wildenstein and Trie‑
benreuth–Tiefenbach slices. Location of the Wildenstein

Sample Alternative Helcionelloida, Pelag‑ Trilobites Member Remarks Collector(s) Coordi‑

ID* sample ID(s) iellida, Stenothecoida, encountered nates
Hyolitha

W2 Hochbehälter, Hyolith genus and species Kingaspidoides Galgenberg Galgenberg Geyer, Kohlmann, N 50° 11′
wasserwerk indet. A frankenwal- Member facies, with Meier, Richter, 47″, E
densis coarse inter‑ Sdzuy 11° 33′
Parasolenopleura calations 29″
wurmi
W8 Str, Weg Helcionella capula Kingaspidoides Wildenstein mostly in Geyer, Meier, N 50° 11′
Helcionella aff. oblonga frankenwal- Member Wildenstein Sdzuy 59″, E
Helcionella sp. A densis facies 11° 33′
“Bemella” sp. A Kingaspidoides 37″
“Igorella” sp. A meieri
Helcionelloid gen. and Parasolenopleura
sp. indet. A wurmi
Latouchella cf. comma
Leptostega frankenwal-
densis
Helcionelloid gen. and
sp. B
Yochelcionella? sp. indet.
Pelagiella sp. A
Stenothecoides? sp. indet.
Grantitheca? klani
Maxilites? sp.
Hyolithid gen. and sp. A
Aladraco schloppensis
Cambrachelous diplo-
prosopus

13

248 G. Geyer et al.

Sample Alternative Helcionelloida, Pelag‑ Trilobites Member Remarks Collector(s) Coordi‑

ID* sample ID(s) iellida, Stenothecoida, encountered nates
Hyolitha

W9 Steinbruch, St, Jincelites cf. vogeli Wildenstein Wildenstein Albert, Geyer, N 50° 12′
Stbr Maxilites? sp. Member facies Klan, Kohlmann, 1″, E 11°
Aladraco schloppensis Meier, Sdzuy, 33′ 35″
Cambrachelous diplo- Richter, Wurm,
prosopus unknown col‑
lector
W11 Galgen-Wiese Cambrachelous diplo- Kingaspidoides Galgenberg Galgenberg Albert, Gandl?, N 50° 12′
prosopus frankenwal- Member facies Geyer, Landing, 13″, E
Hyolith genus and species densis Meier, Richter?, 11° 33′
indet. D Kingaspidoides Sdzuy, Wurm? 44″
laetus?
Acadoparadox-
ides sp. A
Enixus aff.
juvenis
Parasolenopleura
wurmi
W12 Galgen, Helcionella sp. A Kingaspidoides Galgenberg Galgenberg Albert, Gandl, N 50° 12′
Galgen I Grantitheca? klani frankenwal- Member facies Geyer, Hickethier, 15″, E
Hexitheca? sp. densis Hiltl, Kohlmann, 11° 33′
Jincelites cf. vogeli Acadoparadox- Landing, Meier, 45″
Aladraco schloppensis ides? sp. A Paulus, Richter,
Cambrachelous diplo- Parasolenopleura Rückert, Sdzuy,
prosopus wurmi Trapp, Wurm
Hyolithid gen. and sp. B Parasolenopleura
Hyolith gen. and sp. parabolica
indet. E
W12c Galgen II, Aladraco schloppensis Kingaspidoides Wildenstein Galgen II in Sdzuy ca. N 50°
2. Lichtung Nephrotheca? sp. frankenwal- Member Wildenstein 12′ 20″,
densis facies E 11° 33′
46″
W13 Grantitheca? klani Galgenberg Galgenberg Meier ca. N 50°
Member facies 12′ 19″,
E 11° 33′
35″
W13a i, iu Latouchella cf. comma Kingaspidoides Galgenberg partly in Sdzuy N 50° 12′
Grantitheca? klani frankenwal- Member Wildenstein 17″, E
Aladraco schloppensis densis facies 11° 33′
Cambrachelous diplo- Parasolenopleura 32″
prosopus wurmi
Hyolith gen. and sp. Ornamentaspis
indet. C cf. crassilim-
bata
W13b j1, ­j2 Helcionelloid gen. and Wildenstein Wildenstein Sdzuy N 50° 12′
sp. indet. A Member facies 18″, E
11° 33′
33″
W13e H, ­H2 Helcionella capula Kingaspidoides Galgenberg Mostly in Sdzuy ca. N 50°
sp. Member Wildenstein 12′ 20″,
facies E 11° 33′
36″
W14 K2 Hexitheca? sp. Kingaspidoides Galgenberg Galgenberg Albert, Geyer, N 50° 12′
frankenwal- Member facies Hickethier, 28″, E
densis Klan?, Meier, 11° 33′
Parasolenopleura Sdzuy, Tragelehn 51″
wurmi

13
Helcionelloids, stenothecoids and hyoliths from the Cambrian Tannenknock Formation, Germany 249

Sample Alternative Helcionelloida, Pelag‑ Trilobites Member Remarks Collector(s) Coordi‑

ID* sample ID(s) iellida, Stenothecoida, encountered nates
Hyolitha

W14a K1 Grantitheca? klani Kingaspidoides Wildenstein Galgenberg Sdzuy ca. N 50°

Hexitheca? sp. frankenwal- Member facies, partly 12′ 27″,
Jincelites cf. vogeli densis in Wilden‑ E 11° 33′
Hyolith gen. and sp. Parasolenopleura stein facies 50″
indet. E wurmi
Ornamentaspis
sp.
W15a N1 Helcionella sp. A Wildenstein Wildenstein Sdzuy ca. N 50°
Brevitheca? sp. Member facies 12′ 26″,
E 11° 33′
54″
W16b OII Aladraco schloppensis Wildenstein Wildenstein Sdzuy ca. N 50°
Cambrachelous diplo- Member facies 12′ 28″,
prosopus E 11° 33′
56″
W17 OIII Aladraco schloppensis Wildenstein Wildenstein Geyer, Sdzuy ca. N 50°
Member facies 12′ 29″,
E 11° 33′
57″
W18a Q Leptostega frankenwal- Wildenstein Wildenstein Sdzuy ca. N 50°
densis Member facies 12′ 31″,
Pelagiella sp. A E 11° 33′
57″
T2 Grantitheca? klani Kingaspidoides Galgenberg Galgenberg Geyer, Landing, N 50° 11′
Cambrachelous diplo- frankenwal- Member facies Meier 14″, E
prosopus densis 11° 32′
Kingaspidoides 56″
meieri
Parasolenopleura
wurmi
Parasolenopleura
parabolica
Acadoparadox-
ides? sp. A
T2a Grantitheca? klani Kingaspidoides Galgenberg Galgenberg Geyer, Landing, N 50° 11′
Jincelites cf. vogeli frankenwal- Member facies Meier 14″, E
densis 11° 32′
Kingaspidoides 57″
meieri
Parasolenopleura
wurmi
Parasolenopleura
parabolica
T2/T2a Teufelsstein See T2 and T2a See T2 and T2a Galgenberg Galgenberg Albert, Gandl, N 50° 11′
Member facies Hickethier, Sdzuy 14″, E
11° 32′
56″

13

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