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An orthothecid hyolith with a digestive tract from

the early Cambrian of Bornholm, Denmark

Vivianne Berg-Madsen, Martin Valent & Jan Ove R. Ebbestad

To cite this article: Vivianne Berg-Madsen, Martin Valent & Jan Ove R. Ebbestad (2018) An
orthothecid hyolith with a digestive tract from the early Cambrian of Bornholm, Denmark, GFF,
140:1, 25-37, DOI: 10.1080/11035897.2018.1432680

To link to this article: https://doi.org/10.1080/11035897.2018.1432680

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GFF, 2018
VOL. 140, NO. 1, 25–37
https://doi.org/10.1080/11035897.2018.1432680

OPEN ACCESS

An orthothecid hyolith with a digestive tract from the early Cambrian of Bornholm,
Denmark
Vivianne Berg-Madsena, Martin Valentb and Jan Ove R. Ebbestada
a
Museum of Evolution, Palaeontology, Uppsala, Sweden; bDepartment of Palaeontology, National Museum, Cirkusová, Horní Počernice, Czech Republic

ABSTRACT ARTICLE HISTORY

The hyolith assemblage from the early Cambrian of Bornholm, Denmark, shows a higher diversity than Received 7 September 2017
contemporary assemblages in Baltoscandia. The most common species in the Green Shales (Læså Accepted 22 January 2018
Formation, Norretorp Member, Cambrian Stage 3), is Hyolithes [=Hyolithus] (Orthotheca) johnstrupi Holm,
KEYWORDS
1893. A specimen of this species shows a well-preserved and almost complete digestive tract, folded into Orthotheca johnstrupi
an approximately 22 mm long chevron-like structure comprised of at least 20 arcuate loops on the ventral revised; digestive tract;
side and a flattened, gently sinuous to straight anal tube on the dorsal side. The thin, phosphatic outer early Cambrian; Bornholm;
shell layer of the conch is crushed under the digestive tract due to compaction while the digestive tract Denmark
is preserved in three dimensions and appears undisturbed. The shape of the digestive tract is similar to
that of the middle Cambrian Guduguwan hardmani (Etheridge) from Australia and the lower Cambrian
specimens from Russia described by Meškova & Sysoev. The Danish specimen is probably an adult, lending
support to the idea that the orthothecid digestive tract becomes more complex during ontogeny. Hyolithus
(Orthotheca) johnstrupi is revised and here referred to Circotheca Sysoev, 1958.

Introduction Poulsen (1967) described further hyolith species from the

Green Shales and confirmed that Hyolithus (Orthotheca) john-
The hyolith assemblage from the early Cambrian of Bornholm,
strupi was one of the dominant macro-fossils in the siltstone.
Denmark, shows a higher diversity and contains better pre-
Although he no doubt examined the specimen with the digestive
served specimens compared to those of equivalent levels in
tract preserved, it escaped his notice and he stated that “none
Sweden. Three hyolith species from the lower Cambrian Green
were better preserved than those described by Holm, for which
Shales (Læså Formation, Norretorp Member, Cambrian Stage
reason nothing new could be added” (Poulsen 1967, pp. 44–46).
3) of Bornholm, Denmark (Figs. 1 and 2) were included in
The collections at the Geological Museum, Copenhagen, were
the taxonomic study of Swedish Cambrian hyoliths by Holm
examined in connection with the 1999 revision. A specimen, col-
(1893): Hyolithus lenticularis, H. (Orthotheca) johnstrupi and H.
lected at Læså near Vejrmøllegård, which revealed part of a diges-
nathorsti. Re-examination of hyolith material from the Green
tive tract was discovered by J.M. Malinky, who also confirmed
Shales by V. Berg-Madsen confirms the presence of the hyolith-
the identification as H. (Orthotheca) johnstrupi. The specimen
ids Hyolithus nathorsti Johnstrup in Holm, 1893, Lovenedolithes
was collected by K.A. Grönwall in 1892 who noted on the label
groenwalli (Poulsen, 1967) including fragments of helens
that it was a ‘specimen with an unusual structure’. Although this
(see also Martí Mus & Bergström 2007) and an operculum, and
was probably the earliest find of an orthothecid digestive tract
a single specimen of Hexitheca teretiuscula (Linnarsson, 1877).
its significance escaped notice. Thus, the first description of a
Holm (1893) stated that H. (Orthotheca) johnstrupi was by far
fossilized hyolith digestive tract became that of Thoral (1935)
the most common species at the locality near Vejrmøllegård
from the Lower Ordovician of France.
(Fig. 1), and a great number of specimens were found in phos-
Preserved soft parts in hyoliths are rare and only about a
phoritic nodules, including 11 opercula. Through the courtesy
dozen reports of are known with material from the Cambrian,
of Professor J.F. Johnstrup in Copenhagen Holm had had access
Ordovician and Devonian (see Devaere et al. 2014 and Moysiuk
to all this material and also collected in situ himself.
et al. 2017 for references). The soft part morphology is impor-
Although Holm (1893, p. 56) stated most specimens were
tant to distinguish between the order Hyolithida and the order
found in phosphoritic nodules, only one of the here described
Orthothecida. The orthothecid digestive tracts are the best docu-
specimens figured by Holm comes from a nodule, all others
mented and is complex and generally sinuously folded, while it is
from the siltstone proper. Specimens supposedly deposited at the
simple and U-shaped in the hyolithids, which may reflect differ-
Geological Survey of Sweden (SGU) according to Holm (1893,
ent feeding habits. However, Devaere et al. (2014) showed in an
p. 4), where he was employed at that time, have not been located
ontogenetic series of the orthothecid Conotheca subcurvata from
(Malinky & Berg-Madsen 1999).

CONTACT  Vivianne Berg-Madsen  Vivianne.Berg-Madsen@em.uu.se

© 2018 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creativecommons.org/licenses/by-nc-nd/4.0/),
which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or built upon in any way.

Published online 06 Apr 2018

26   V. BERG-MADSEN ET AL.

(A) (B) (C)

(D)

Figure 1.  A. Maps showing the position of Bornholm relative to the Danish mainland. B. The island with major roads and communities. C. Area with the locality
Vejrmøllegård at Læså rivulet near Aakirkeby and Grødby rivulet, exact collecting locality unknown. D. Geological map of southern Bornholm showing sedimentary units
and the position of Vejrmøllegård relative to the Læså rivulet and Læså Formation. Geological map based on Graversen (2009).

the Cambrian of France that the digestive tract is U-shaped in
the earliest growth stages and becomes sinuous only in the adult
stage. They suggested that this could imply a distinct phyloge-
netic relationship between the two hyolith orders, and further
speculated that the change in the morphology in the orthothe-
cid digestive tract reflected a change in feeding habit during the
growth of the animal.
A description of the digestive tract from the lower Cambrian
of Bornholm is presented here, but our study also incited a rede-
scription of Hyolithus (Orthotheca) johnstrupi Holm, 1893 and
a reassignment of the species to Circotheca Sysoev, 1958 emend.
Berg-Madsen & Malinky 1999 which will be used throughout
the text.
Geological setting
The geology of southern Bornholm is characterized by a number
of fault blocks (Fig. 1) and exposures are primarily found only
along rivulets or along the coast (Hansen 1936; Poulsen 1967).
Originally, the lower Cambrian strata were regarded as one uni-
form unfossiliferous quartzite and very little attention was paid to
the later called Green Shales (Pingel 1828; Forchhammer 1835).
Eventually, J.F. Johnstrup and his students collected fossils from
the strata from 1869 onwards, and Johnstrup was the first to sub-
divide the lower Cambrian on Bornholm (Johnstrup 1874, 1891).
The lower Cambrian of Scandinavia was revised by Nielsen &
Schovsbo (2007), which is now divided in ascending order the
Hardeberg Formation overlain by the Læså Formation, which
includes the Norretorp Member overlain by the Rispebjerg
Member (Fig. 2). The Norretorp Member, previously known as
the Green Shales, comprises about 100 m of glauconitic siltstone
and storm generated sand beds (Nielsen & Schovsbo 2011). The
siltstone is thinly bedded, bioturbated and the surface normally
of rusty brown colour due to the weathering of the glauconite
and pyrite; the fresh colour is greenish-grey. Levels bearing phos-
phoritic nodules are found throughout the stratum and hyolith-
ids and orthothecids have been reported from both the nodules
and surrounding siltstone. Poulsen (1967), and before him both

Figure 2. Correlation between Siberian and Baltoscandian stages, acritarch zones
and lithostratigraphic units on Bornholm. Modified from Moczydłowska & Vidal
(1992) and Nielsen & Schovsbo (2011).
Johnstrup (1891) and Grönwall (1899), noted the presence of
specimens in a “grey limestone” within the Green Shales. This
unit was later correctly identified as a thin bed of fine grained,
very dense sandstone, at places with calcitic matrix, in the lower
part of the Green Shales at Læså and Grødbyå (Hansen 1936).
Hansen (1936) recognized levels especially rich in hyo-
liths and phosphoritic nodules in the Læså stream south-west
of Vejrmøllegård, belonging to the upper middle part of the
member. For details of localities and maps see Hansen (1936,
pp. 50–52, Fig. 8); subsequently used by Poulsen (1967) and
Moczydłowska & Vidal (1992).
The Norretorp Member represents an offshore low-energy
shelf environment and fairly deep water (Moczydłowska & Vidal
1992, Fig. 11). The siltstone sequence represents a shallow marine
shelf and progressing coastline following transgressive and
regressive episodes. Occasional fluxes of more coarse-grained
storm deposits occur throughout the sequence. Judged from the
weathered surface of the samples the hyoliths most often lie par-
allel to the bedding plane but not in living position.
The lower Cambrian strata on Bornholm are equivalents of
the Siberian Nemakit-Daldynian to Atdabanian Stages (Mens
et al. 1987, 1990). The Læså Formation was deposited during
Cambrian Series 2 (Stage 3) (Fig. 2), and correlates with the
Atdabanian Stage of Siberia. The lower part of the formation
belongs to the Skiagia ornata-Fimbriaglomerella membranacea
Acritarch Zone and the upper part the Heliosphaeridium dis-
similar-Skiagia ciliosa Acritarch Zone (Moczydłowska & Vidal
1992). Specimens of Circotheca johnstrupi are found at several
localities representing the lower and middle part of the Norretorp
Member (Poulsen 1967; Berg-Madsen pers. observation). Thus,
the Bornholm specimen with the preserved digestive tract is of
the same age as the specimens with digestive tracts described
from the Atdabanian of Siberia (Meškova & Sysoev 1981). The
GFF   27
Bornholm specimen is younger than hyoliths with preserved
digestive tracts from the Terranuevian of Montagne Noire
(Devaere et al. 2014).
The digestive tract
Hyolithids with soft parts preserved are generally rare but
Moysiuk et al. (2017) reported more than 200 hyolithid spec-
imens of Haplophrentis Babcock & Robison, 1988 from the
Burgess Shale and Spence Shale Lagerstätten in western North
America with preserved soft tissues. However, the digestive tract
from Circotheca johnstrupi is only the third occurrence from the
early Cambrian to be reported. Meškova & Sysoev 1981 isolated
two fragments of digestive tracts from the early Cambrian of
Siberia. These were extracted from the matrix by acetic acid and
although associated with several hyolithids and orthothecids,
were not referable to any genera or species. Recently, Devaere
et al. (2014) recorded a substantial number of specimens of the
species Conotheca subcurvata Yu, 1974 from the early Cambrian
(Terraneuvian) of Montagne Noire, France with digestive
tracts preserved. Several more orthothecids species, of middle
Cambrian to Lower Devonian age, with preserved soft parts have
been recorded (see Devaere et al. 2014 for references).
The most complete specimen of Circotheca johnstrupi is
MGUH 31848 (Fig. 3), but only about half the length of the
specimen was visible when discovered. Careful preparation by V.
Berg-Madsen subsequently revealed both the entire apical part
and the operculum. It was not attempted to free more of the ada-
pertural part of the conch as enough of the aperture was visible.
The conch measures 42 mm in length from the tip of the apex to
the rim of the aperture and has an apertural diameter of about
7 mm (Fig. 3A). At least eight septa are present within the first
adapical 10 mm, followed by 12 mm of ‘empty’ space now partly
filled with fragments of crushed shell and sediment grains (Fig.
3A, B and H). The convoluted digestive tract is preserved in partly
phosphatized sediment, and folded into a chevron-like structure.
It is about 22 mm long with a posterior width of 1.5 mm, the
anterior width is 6 mm and the cross section is elliptical. Both the
anterior and posterior parts are slightly damaged and obscured
by microbial growth and consequently, the exact length cannot
be measured (Fig. 3B, F and G). The position of the digestive
tract is slightly skewed from that of the operculum as judged
from the axis of the conch, and with the ventral side facing the
flattened venter of the conch. On the ventral side, a series of at
least 20 arcuate loops can be counted, while on the dorsal side
the central part is a 0.9 mm wide, flattened (empty?) and gen-
tly sinuous (distortion?) straight anal tube with the posterior
u-shaped bend missing (Fig. 3B, G). A gap of 1.5 mm separates
the digestive tract from the in situ operculum preserved as a low
eccentric cone. Only the exterior of the operculum is preserved
and sediment fills the anterior-most part of the conch.
The thin, phosphatic outer shell layer is crushed under the
digestive tract due to compaction but the digestive tract itself
shows no sign of compaction, suggesting that phosphatization
was finished long before the shell was compacted.
The digestive tract, preserved in three dimensions, was
removed from the conch almost completely (broke in two pieces)
and only the posterior 5 mm were impossible to extract without
damage. Once the digestive tract was removed its ventral side
28   V. BERG-MADSEN ET AL.

Figure 3. Circotheca johnstrupi (Holm, 1893). Specimen MGUH 31848 with preserved digestive tract. A. External mould with digestive tract removed. B. Specimen
with digestive tract and anal tube in place. C. Latex cast of external mould. D. External mould showing external surface of operculum. E. Latex cast of external mould of
operculum showing external surface of operculum. F. Ventral side of extracted digestive tract. G. Dorsal side of extracted digestive tract and anal tube. H. Posterior end
of conch with septa. I. Detail of digestive tract in left ventral oblique view. J. Detail of digestive tract in right lateral view. Læså Formation at Vejrmøllegård, Læså rivulet.
Collected by K.A. Grönwall. Scale bars = 0.1 cm.
 GFF   29

could be examined, and it also revealed the previously partly

obscured operculum and ventral side of the conch, also allowing
a latex cast to be made (Fig. 3C, E).
The shape of the digestive tract of Circotheca johnstrupi is
closer to that of Guduguwan hardmani (Etheridge in Foord, 1890)
from the middle Cambrian of the Northern Territory in Australia
(Kruse 1997) than those known from the early Cambrian of
Montagne Noire, France (Devaere et al. 2014, Fig. 10). However,
its morphology is similar to that described by Meškova & Sysoev
(1981) from the early Cambrian specimens of Siberia.
Devaere et al. (2014) suggested that a simple U-shaped diges-
tive tract represents an early stage of development in orthothe-
cid hyoliths, the folding advancing at later growth stages of the
animal. All specimens of Circotheca johnstrupi from the lower
Cambrian of Bornholm are remarkably uniform in size, and very
few are so small that they can be regarded as juvenile. The spec-
imen with the digestive tract is most likely an adult individual,
which lend support to the suggestion by Devaere et al. (2014).
Although without early stages with a preserved digestive tract,
we can only see the final development.
The discussion of dorsal/ventral position in hyoliths is closely
related to the mode of life and in several hypotheses, among oth-
ers discussed by Yochelson (1961), Fisher (1962), Marek (1963),
Runnegar et al. (1975), Marek & Yochelson (1976), Sysoev (1973,
1976, 1984), Dzik (1980), Kruse (1997), and Marek et al. (1997).
The latter authors regarded the inferred dorsal/ventral position
and orientation of the gut in the single Australian specimen
described by Kruse (1997), where the flat side was interpreted
as dorsal, as a ‘preservational oddity’. The view of Kruse (1997)
has not since been endorsed and the inflated part of the conch in
both hyolithids and orthothecids has been considered as dorsal.
In a conch with a nearly circular cross section, the maximum
width of the digestive tract must be less than the diameter of
the conch. Once free of assumed organic filaments keeping it
in place, the intestine has a much greater chance to move in
a nearly circular conch if this is being turned upside down or
rolled. This means it can end up in any position relative to the
operculum. The conch coquina on the bedding plane shown
by Kruse (1997, Fig. 2A) indicates a high energy environment.
Thus, it is likely that tumbling of the intestine took place, which
explains its upside down orientation.

Systematic palaeontology
Remarks. – In a footnote Holm (1893, p. 7) explains the reason
for spelling the genus Hyolithus rather than the nowadays still
widely used Hyolithes introduced by Eichwald (1840), pointing
out that it was entymologically incorrect, and he would use ‘the
only correct form of the name Hyolithus’ (translated here). That
the latter term is still used, as exemplified for instance by the Figure 4.  Circotheca johnstrupi (Holm, 1893). Lectotype MGUH 1506. A. Partial
synonymy list presented below, could be because the footnote steinkern and partial external mould in right oblique view. B. Reproduction
of original figure in Holm (1893, pl. 1, Fig. 28). C. Right lateral view of latex cast
may have escaped notice. Holm’s work was as well written in of external mould. D. Detail of ornamentation. Line across the conch may mark
Swedish which could also be a hinder for the understanding by possible shell breakage. Læså Formation at Vejrmøllegård, Læså rivulet. Collected
non-Scandinavian readers. by J.F. Johnstrup. Scale bars in A–C = 0.5 cm. Scale bar in D = 0.1 cm.
For the transliteration of Russian names (for example Syssoiev;
Sysoev; Sysoyev) and references, we use the ISO 9:1995 translit- Family Circothecidae Missarževskij 1969, emended Berg-
eration of the Cyrillic characters (i.e. Sysoev). Madsen & Malinky 1999
Class Hyolitha Marek 1963 Genus Circotheca Sysoev, 1958, emended Berg-Madsen &
Order Orthothecida Marek 1966 Malinky, 1999
30   V. BERG-MADSEN ET AL.
Figure 5.  Circotheca johnstrupi (Holm, 1893). A. Paralectotype MGUH 1507,
partial steinkern, left lateral view. White infilling on either side is old ‘latex’. B.
Reproduction of original figure in Holm (1893, pl. 1, Fig. 30). C. Paralectotype
MGUH 1514, partial steinkern in dorsal view with phosphatized bacterial matter
along the centre and septa at the apex. D. Reproduction of original figure in Holm
(1893, pl. 1, Fig. 71). Læså Formation at Vejrmøllegård, Læså rivulet. Collected by J.F.
Johnstrup. Scale bars = 0.5 cm.
Type species. – By original designation of Sysoev (1958), Hyolithus
(Orthotheca) stylus Holm, 1893, p. 52, pl. 1, Figs. 16–20; pl. 6,
Figs. 6–9, from the mid-Cambrian Series 3 part of the Alum
Shale Formation (Paradoxides paradoxissimus Super Zone,
Lejopyge laevigata Zone) in Västergötland, Sweden.
Circotheca johnstrupi (Holm, 1893) emended
Figs. 3–11
1893 Hyolithus (Orthotheca) johnstrupi n. sp. – Holm, p. 56, pl.
1, Figs. 28–32, 71.
Non 1893 Hyolithus (Orthotheca) johnstrupi n. sp. – Holm,
p. 56, pl. 1, Fig. 33.
1902 Hyolithus johnstrupi Holm – Grönwall, p. 18, Fig. 4.
1930 Hyolithus johnstrupi Holm – Milthers p. 33, Fig. 9.
Reproduced from Holm.
1938 Hyolithes johnstrupi Holm – Rosenkrantz, p. 5, pl. 2, Fig. 1.
1946 Hyolithes (Orthotheca) johnstrupi Holm – Sinclair, p. 77.
1962 Orthotheca johnstrupi Holm – Sysoev, p. 10.
1966 Hyolithes johnstrupi Holm – Rasmussen, p. 27, text-Fig. 2
(pars)., left image only.
1967 Hyolithes johnstrupi Holm – Schiønning & Wagner, p. 66,
Fig. 58. Redrawn from Grönwall.
1967 Orthotheca johnstrupi Holm – Poulsen, p. 21.
Non 1988 Hyolithes (Orthotheca) johnstrupi Holm – Landing,
p. 675.
Material. – Lectotype, here designated, MGUH 1506 (Holm
1893, pl. 1, Figs. 28–29; here Fig. 4). Paralectotypes MGUH 1507
(Holm 1893, pl. 1, Fig. 30; here Fig. 5A), MGUH 1509 (Holm
1893, pl. 1, Fig. 32; here Fig. 6E, F) from Læså, Bornholm, col-
lected by Johnstrup. MGUH 1514 (Holm 1893, pl. 1, Fig. 31;
here Figs. 5C and 11D). Paralectotype? MGUH 1508 (Fig. 6A,
B). Additionally, figured MGUH specimens are MGUH 31848–
MGUH 31857. All specimens are housed in the Natural History
Museum of Denmark, University of Copenhagen (MGUH).
Stratigraphy. – Læså Formation, Norretorp Member, upper
Skiagia ornata–Fimbriaglomerella membranacea to lower
Heliosphaeridium dissimilare–Skiagia ciliosa acritarch zones.
Cambrian Series 2 (Stage 3).
Diagnosis. – Circotheca having ventral apertural edge with short,
ligula-like projection; shell with transverse rugae only and with
subtriangular cross section with highly rounded edges; exterior
of operculum with prominent concentric rugae and interior with
finer rugae.
Description. – Orthoconic conch with small apical angle and
slender appearance; cross section round at apex growing increas-
ingly subtriangular with highly rounded edges; venter flat and
grading through rounded lateral edges into inflated dorsum that
lacks a clearly defined median ridge. Aperture almost planar with
a faint ligula-like projection along ventral rim. Surface of shell,
at least on part of dorsal and part of lateral margins, covered
with low transverse rugae but with regions of irregular width
and of irregular spacing containing growth lines; on venter lines
and rugae are convex towards aperture to follow edge of ventral
ligula-like projection. At least eight septa present in apical region;
internal mould smooth.
Operculum heavily rounded subtriangular in outline. Conical
shield broad and flat with very gentle fold in central portion;
 GFF   31

Figure 6. Circotheca johnstrupi (Holm, 1893). A, B. Steinkern of MGUH 1508 purportedly the original of Holm (1893, pl. 1, Fig. 31). C, D. Steinkern of MGUH 31849
proposed here as true original of Holm (1893, pl. 1, 31). E, F. Steinkern of MGUH 1509, original of Holm (1893, pl. 1, Fig. 32). Læså Formation at Vejrmøllegård, Læså rivulet.
Collected by J.F. Johnstrup. Scale bars in A, B, D = 0.5 cm. Scale bars in C, E = 0.1 cm.

surface of cardinal shield rounded and slightly convex with con- the fold in the growth lines and rugae on the venter (Fig. 9), and
stant slope throughout; transition between conical and cardinal from the fact that the operculum is not planar, with the cardi-
shields forms an angle of about 8 degrees. Exterior with con- nal and main shields intersecting at an oblique angle. Circotheca
centric rugae but fainter and fewer in number than on interior. johnstupi differs from the mid-Cambrian C. smetanai Valent et
al., 2012 (Barrandian Area, Czech Republic) in the conch sculp-
Remarks. – This species was originally based on three internal
ture where C. smetanai has very narrow longitudinal ribs and
moulds of conchs (MGUH 1506, MGUH 1507, MGUH 1514)
in the cross section where C. smetanai shows a circular cross
and three opercula (MGUH 1508–MGUH 1510) (Holm 1893, pl.
section. It should be noted that the cross section of C. johnstrupi
1, Figs. 28–33, 71). MGUH 1506 is selected as the lectotype (Fig.
(Holm, 1893, pl. 1, Fig. 29) represents the posterior subcircular
4). It is preserved as an internal and external mould which meas-
part of the internal mould of the conch, which otherwise gradu-
ure 40 mm long, and has an apertural width of 5 mm. Additional
ally becomes more subtriangular anteriorly as seen by the shape
specimens furnish details of the shell, the digestive tract and in
of the opercula (Fig. 6).
particular of the operculum (Fig. 8).
Holm (1893) noted that this species was similar to Hyolithus
The species is assigned to Circotheca Sysoev, 1958 under the
(Orthotheca) degeeri Holm, 1893, but types of the latter are
revised concept of this genus given by Berg-Madsen & Malinky
incompletely preserved, and indeed, Holm himself even sug-
(1999). This species differs from the type species C. styla, in hav-
gested that better preserved specimens of that species may show
ing transverse ornament only, and from both the Ordovician
it to be the same as Circotheca johnstrupi. Ironically, it is the name
C. caperai Marek, 1983 from Montagne Noire, France, and C.
of the former that has come to be widely used for conical fossils
neptis Marek, 1989 from the Barrandian Area, Czech Republic,
in the Cambrian (see Malinky & Berg-Madsen 1999, pp. 54, 55
by having a ventral ligula-like apertural projection. Although
for details of ‘O.’ degeeri), whereas it is the later species that is far
the aperture on all specimens of C. johnstrupi is incomplete, the
better preserved. Cobbold (1919) noted that the cross section of
presence of the projection is certain, having been inferred from
32   V. BERG-MADSEN ET AL.
Figure 7. Orthothecid? MGUH 1510. A. Internal surface of supposed operculum.
B. Enlarged reproduction of original figure in Holm (1893, pl. 1, Fig. 33). Læså
Formation at Vejrmøllegård, Læså rivulet. Collected by J.F. Johnstrup. Scale
bar = 0.1 cm.
H. (O.) degeeri is similar to that of H. (O.) johnstrupi, and he later
figured (Cobbold 1931, pl. 41, Fig. 5) an operculum Hyolithes
(Orthotheca) sp. indet. (MGUH 31857), noting similarities with
those of C. johnstrupi figured by Holm. Whereas C. johnstrupi
is founded on a sound morphological basis with features sup-
porting placement under Circotheca, Malinky & Berg-Madsen
(1999) judged ‘O.’ degeeri to be unrecognizable.
Holm (1893) included both Circotheca. johnstrupi and H. (O.)
degeeri under the division Complanati, in which the ventral side is
slightly flattened. The incomplete preservation of the latter species
precludes further comparison (Malinky & Berg-Madsen 1999).
Holm (1893, pl. 1, Fig. 71) also noted that the conch (i.e.
MGUH 1514, Fig. 5C, D) may have as many as seven septa, but
because of preservation, the exact number may be difficult to
determine. Several other specimens of Circotheca johnstrupi also
reveal two or more septa of which the best preserved are those
in MGUH 31848 (Fig. 3A, H).
Circotheca johnstrupi is a definitive representative of
Circotheca from the lower Cambrian. The name Circotheca has
been widely used for tubular fossils from this level, particularly
in the Tommotian of Siberia and Meischucunian of China (Qian
& Bengtson 1989; Malinky & Berg-Madsen 1999 and references
therein). Many of these fossils are poorly preserved moulds and
casts of uncertain affinity. Use of the name Circotheca for any of
them distorts the geographical and stratigraphical distribution
of that genus. However, C. johnstrupi (Holm, 1893) demonstrates
that authentic Circotheca has a range of at least lower to middle
Cambrian.
Circotheca johnstrupi (Holm, 1893) is by far the most common
hyolith in the Norretorp Member but has so far not been found
elsewhere in Scandinavia. The species is found at several locali-
ties representing the lower and middle part of the member. The
specimens are preserved either as internal moulds, partly empty
conchs or as external mould of the conchs. The conchs often con-
tain single strands or sheets of microbial growth which eventually
develop to solid phosphate filling the conch completely, as also
demonstrated by Devaere et al. (2014).
Reconstructions after Holm’s illustrations of Circotheca john-
strupi have frequently been published, but most authors did not
distinguish between hyolithids and orthothecids (see synonymy
list). A reconstruction showing Hyolithes johnstrupi with helens
(exclusively found in hyolithids) was published by Rasmussen
(1966), which since then has been reproduced in textbooks, pop-
ular literature and recently on internet at least 10 times in most
cases as Hyolithes or a hyolithid (see for instance Rasmussen
1969, p. 290; Poulsen & Poulsen 1975, p. 57, text-Fig. 43). We
refrain from listing all these publications, except for those works
where the reconstruction is illustrated next to a photograph of
the lectotype MGUH 1507.
Opercula
Most opercula have a height between 4 and 6 mm and a width
between 5 and 7  mm. Almost all are found well inside the
conch and usually with the concave side lying against the inte-
rior of the shell, and the convex side pointing towards infill-
ing of the conch (Figs. 6 and 8B–D, F–H). The operculum
in Fig. 10B is preserved with the apex pointing towards the
interior of the shell. Holm (1893, p. 57) mentioned the simi-
larity of these opercula with the “brachiopods Acrothele and
Discinella” (=Mobergella but referred to Discinella by Moberg
(1892)). From the figures (Holm 1893, pl. 1, Figs. 31, 32) the
shape seems close to that of the lingulate brachiopod Acrothele
whereas neither shape, ornament nor interior are close to that
of the enigmatic fossil Mobergella.
Holm (1893, pl. 1, Figs. 31, 32) figured two opercula taken
out of their context as the position inside the conch is omitted
(Fig. 6). We have some doubt that the operculum MGUH 1508
labelled as part of the type series is the one figured in Holm
(1893, pl. 1, Fig. 31). It is slightly chipped, positioned so that
it is difficult to angle for a drawing and it is difficult to see any
details without colouring it dark and whitening with ammonium
chloride sublimate (Fig. 6A, B). Another specimen available to
Holm, but not labelled as part of the type series, MGUH 31852
(Fig. 6C, D), seems to be a better match as it is perfectly preserved
and details would be easy to delineate in a drawing. The size of
the two afore-mentioned specimens is about equal and matches
that shown by Holm (1893).
The only operculum showing the interior (MGUH 31850,
Figs. 8A, E and 11A) is free of overlying sediment, the exterior
 GFF   33

Figure 8. Circotheca johnstrupi (Holm, 1893). A, E. MGUH 31850. External mould with bacterial mat near operculum and internal surface of operculum. B, F. MGUH
31851. External mould with impressions of ornamentation and external surface of operculum. C, D, G. MGUH 31852. External mould, latex cast of external mould and
external surface of operculum. H. MGUH 31853. Operculum showing external surface. Læså Formation at Vejrmøllegård, Læså rivulet. Collected by J.F. Johnstrup. Scale
bars in = 0.5 cm.

side obscured by microbial growth. The operculum is almost
circular in outline with a broad and flat main shield with a
very shallow fold in the central portion, eccentric apex and a
rounded cardinal shield, the shields intersecting at an oblique
angle. It has a maximum width and height of 4.9 and 4.7 mm,
respectively. It is notable that the operculum lacks internal
characters like clavicles and cardinal processes. The conch is
incomplete and it is difficult to make out whether or not the
operculum is withdrawn, but a slightly skewed position may
be possible.
The record of this species allows for additional documentation
of the fact that the orthothecid operculum was narrower than
the aperture, and therefore could be withdrawn into the conch
as suggested by Marek (1966, Fig. p. 91; 1967) and illustrated by
Devaere et al. (2014, Fig. 4I, K, p. 5). We exclude the specimen
MGUH 1510 described by Holm (1893, pl. 1, Fig. 33, here Fig.
7) from the genus Circotheca. Holm illustrated what he believed
to be the interior of the operculum possessing a deep, oval
depression surrounded by cardinal processes. It cannot be com-
pletely ruled out that the structures are malformations caused
by secondary growth. The usual morphology of the interior of
the operculum of representatives of the genus Circotheca shows
distinct paired bilobate cardinal processes (see type species
Circotheca styla (Holm, 1893) in Berg-Madsen & Malinky 1999
(text-Fig. 9E, F and R) or Circotheca smetanai Valent et al. 2012
(Fig. 2A, B)) which are not present in any form at Holm’s speci-
men. Although conchs are present in the same sample, the oper-
culum is not connected with any of these. If indeed an operculum
and not a piece of isolated shell, the possibility exists that it is an
operculum belonging to one of the hyolithid species (for instance
Hyolithus nathorsti Johnstrup in Holm) present in the Norretorp
Member, the interior of which are equally poorly known (Holm
1893; Poulsen 1967).
The apparent lack of cardinal processes on the internal sur-
face of the operculum of Circotheca johnstrupi may suggest that
it was fastened in the epidermis much like in gastropods. This
may also explain why almost all opercula referable to Circotheca
johnstrupi are found inside the conch, suggesting that they were
pulled in by the visceral mass. Sediment infilling the apertural
part of the conch would prevent the operculum from disap-
pearing. Sediment infill could also explain why some opercula
were kept in place allowing development of microbial growth
34   V. BERG-MADSEN ET AL.
Figure 9. Circotheca johnstrupi (Holm, 1893). A, B. MGUH 31854, latex cast of ventral side of external mould showing well-preserved ornamentation. Læså Formation at
Grødby rivulet. Collected by J.F. Johnstrup. C, D. MGUH 31855. Dorsal, left lateral and ventral views of latex cast of external mould showing well-preserved ornamentation.
Scale bar in A = 0.5 cm. Scale bars in B–E = 0.1 cm.
in the reducing conditions inside the conch developed during
decomposition.
Shell
Already Holm (1893, p. 1) noted three to four layers in the
shell, assuming it consisted of calcium carbonate; the calcium
carbonate has secondarily been replaced by calcium phosphate.
He also believed the conch and the operculum consisted of two
different types of material and compared this with the situation
in certain gastropods corneous opercula. In several conchs, two
and perhaps three layers can be distinguished (MGUH 31854,
Fig. 10D; MGUH 1514, Fig. 11B). The variable expression and
numbers of rugae on the exterior relative to the interior of the
operculum shows that there are distinct differences in the respec-
tive shell layers. The present material is too scarce and badly
preserved to provide further information.
Microbial growth
Phosphatized microbial growth is often found around the oper-
cula, and in some cases the internal mould seems to consist of
more or less dense filaments of phosphatized bacteria (Fig. 11).
Holm (1893) described this infill as being like cinder (Holm
1893, pl. 1, Fig. 71; here Figs. 5C, D and 11B). Microbial growth
is represented by botryoidal, uniseriate and branching filaments
as well as diagenetic overgrowth, especially below the only oper-
culum revealing the interior the filaments form film or sheets
MGUH 31850 (Fig. 11A). Some of these filamentous structures
may reflect direct phosphatization of the partly decomposed soft
tissue (see discussion in Zhao & Bengtson 1999; Devaere et al.
2014). The posterior septated part of the conch is always devoid
of microbial growth.
Summary
The digestive tract documented in the Cambrian orthothecid
herein is the first such from Denmark (and Scandinavia), and
only the third known with certainty from the lower Cambrian.
It is preserved in three dimensions and can be mechanically
detached from the conch allowing inspection of all sides. The
importance of such a specimen in understanding hyolith pal-
aeobiology and palaeoecology cannot be overstated, given that
few other such specimens have been found. Its significance may
be judged in part from the general reconstruction of hyolith soft
part morphology given by Runnegar et al. (1975), who conferred
separate phylum status on the hyoliths because of presumed dif-
ferences between them and the Mollusca, where hyoliths had
been traditionally placed (Marek & Yochelson 1964, 1976).
The systematic position of hyoliths was summarized and dis-
cussed by Malinky & Yochelson (2007) who regarded these as a
class within the molluscs. Sun et al. (2016) mentioned the phy-
logenetic relationships of molluscs, sipunculans, hyolithids and
orthothecids based mainly on the first appearance of group and
development of the gut and intestine and orthothecids are treated
here as a more primitive/ancestral group of hyoliths. Based on
excellent material with preserved soft tissues of the hyolithid
Haplophrentis Babcock & Robison 1988 from the Burgess Shale,
Canada and the Spence Shale, USA, Lagerstätten, Moysiuk et al.
(2017) published a new hypothesis that hyoliths can be placed

Figure 10. Circotheca johnstrupi (Holm, 1893). A, B. MGUH 31856. Steinkern of
conch in ventral and left lateral views with operculum resting on floor of conch
with external side down. C. MGUH 31857. Ventral side of nearly complete conch. D.
MGUH 31854. Detail of latex cast of conch showing two shell layers. See Fig. 9A for
entire specimen. Scale bars in A, C = 0.5 cm. Scale bars in B, D = 0.1 cm.
among the Lophophorata. Based on their findings, this affinity
seems to be clear for the hyolithids but such high taxonomic
placement for orthothecids is not at the moment possible as the
organization of the internal organs remains unknown.
Marek et al. (1997) used the rare preservation of the intes-
tines in orthothecids for a palaeoecological interpretation. They
naturally reasoned that in order for the orthothecid digestive
GFF   35
Figure 11.  Circotheca johnstrupi (Holm, 1893). A. MGUH 31850. Detail of
phosphatized bacterial mesh below operculum. See Fig. 8A for entire specimen. B.
MGUH 1514. Detail of posterior part of conch with phosphatized bacterial infilling.
Numbers 1–3 indicate three possible shell layers. See Fig. 5C for entire specimen.
Scale bars in = 0.1 cm.
tract to be preserved, it must filled with sediment; and the fact
that intestines are in some cases preserved fully three-dimen-
sional supports the notion that these animals were deposit
feeders ingesting sediment. In contrast, experimental evidence
showed hyolithids were suspension feeders. Lastly, the remains of
microbial growth beneath the operculum and within the conch
demonstrate the early microbially mediated phosphatization of
this material which was also shown in the early Cambrian mate-
rial from France described by Devaere et al. (2014).
Acknowledgement
J.M. Malinky is thanked for his initial enthusiastic involvement in this
study about 20  years ago. Original specimens of Holm and subsequent
material from the Norretorp Member, Bornholm, were kindly made avail-
able from the Geological Museum in Copenhagen and museum numbers
were provided by A.R. Bashforth. M. Valent was financially supported by
36   V. BERG-MADSEN ET AL.
the Ministry of Culture of the Czech Republic (DKRVO 2018/06, National
Museum, 00023272 – M. Valent). The manuscript was improved consider-
ably by comments from two reviewers and robust feedback from the editor.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported by Ministerstvo Kultury [grant number DKRVO
2018/06].
References
Babcock, L. & Robison, R.A., 1988: Taxonomy and paleobiology of some
Middle Cambrian Scenella (Cnidaria) and hyolithids (Mollusca)
from western North America. University of Kansas Paleontological
Contributions 121, 1–22.
Berg-Madsen, V. & Malinky, J.M., 1999: A revision of Holm’s late Mid and
Late Cambrian hyoliths of Sweden. Palaeontology 42, 841–885.
Cobbold, E.S., 1919: Cambrian Hyolithidae, etc. from Hartshill in the
Nuneaton District, Warwickshire. Geological Magazine 6, 149–158.
Cobbold, E.S., 1931: Additional fossils from the Cambrian rocks of
Comley, Shropshire. Quarterly Journal of the Geological Society, London
87, 459–512.
Devaere, L., Clausen, S., Alvaro, J.J., Peel, J.S. & Vachard, D., 2014:
Terreneuvian Orthothecid (Hyolitha) Digestive Tracts from
Northern Montagne Noire, France; Taphonomic, Ontogenetic
and Phylogenetic Implications. PLoS One 9(2), e88583. doi:
https://doi.org/10.1371/journal.pone0088583.
Dzik, J., 1980: Ontogeny of Bactrotheca and related hyoliths. Geologiska
Föreningens i Stockholm Förhandlingar 102, 223–233.
Eichwald, K.E. von, 1840: Über das silurische Schichtensystem in Esthland.
Zeitschrift für Natur- und Heilkunde der königlische Medicinisch-
chirurgische Akademie St. Petersburg 1/2, 1–210.
Fisher, R.W., 1962: Small conoidal shells of uncertain affinities. In R.C.
Moore (ed.): Treatise on invertebrate paleontology. Part W. Miscellanea,
W98–W143. Lawrence, KS, Geological Society of America and
University of Kansas Press.
Foord, A.H., 1890: Descriptions of fossils from the Kimberley district,
Western Australia. Geological Magazine 7(98–106), 145–155.
Forchhammer, G., 1835: Danmarks geognostiske forhold. Indbydelsesskrift
til Reformationsfesten den 14’de November 1835, 122. Kjøbenhavn, J.H.
Schultz.
Graversen, O., 2009: Structural analysis of superposed fault systems of
the Bornholm horst block, Tornquist Zone, Denmark. Bulletin of the
Geological Society of Denmark 57, 25–49.
Grönwall, K.A., 1899: Bemærkninger om de sedimentære dannelser
på Bornholm og deres tektoniske forhold. Danmarks geologiske
Undersøgelse, 2 Række 10, 1–52.
Grönwall, K.A., 1902: Grundrids af Bornholms geologi, 36. Rønne, Frits
Sørensen.
Hansen, K., 1936: Die Gesteine des Unterkambriums von Bornholm.
Danmarks geologiske Undersøgelse, 2 Række 62, 1–194.
Holm, G., 1893: Sveriges Kambrisk-Siluriska Hyolithidæ och Conulariidæ.
Sveriges geologiska undersökning C 112, 1–172.
International Standard ISO 9:1995: Information and documentation –
transliteration of Cyrillic characters into Latin characters – Slavic and
non-Slavic languages. ICS 01.140.10. International Organization for
Standardization, Genève.
Johnstrup, J.F., 1874: Oversigt over de palæozoiske dannelser på Bornholm.
Beretning 11te skandinaviske naturforskermøde i Kjøbenhavn 1873, 10 pp.
Johnstrup, J.F., 1891: Abriss der Geologie von Bornholm, als Führer zu der
Exkursion der Deutschen Geologische Gesellschaft der Insel Bornholm
im Anschluss an die allgemeine Versammlung in Greifswald 1889, 66.
Greifswald, Deutsche geologische Gesellschaft.
Kruse, P.D., 1997: Hyolith guts in the Cambrian of northern Australia –
turning hyolithomorphs upside down. Lethaia 29, 213–217.
Landing, E., 1988: Lower Cambrian of eastern Massachusetts: stratigraphy
and small shelly fossils. Journal of Paleontology 62, 661–695.
Linnarsson, G., 1877: Fynd av Andrarumskalk på Hunneberg i Västergötland.
Geologiska Föreningens i Stockholm Förhandlingar 3, 346–347.
Malinky, J.M. & Berg-Madsen, V., 1999: A revision of Holm’s early and
early mid Cambrian hyoliths of Sweden. Palaeontology 42, 25–65.
Malinky, J.M. & Yochelson, E.L., 2007: On the systematic position of
the Hyolitha (Kingdom Animalia). Memoirs of the Association of
Australasian Palaeontologists 34, 521–536.
Marek, L., 1963: New knowledge on the morphology of Hyolithes. Sborník
geologických ved, rada Paleontologie 1, 53–72.
Marek, L., 1966: New hyolithid genera from the Ordovician of Bohemia.
Casopis Národního Muzea 135, 89–92.
Marek, L., 1967: The class Hyolitha in the Caradoc of Bohemia. Sborník
geologických ved, rada Paleontologie 9, 51–113.
Marek, L., 1983: The hyoliths in the Arenigian of Montagne Noire. In
R. Courtessole, L. Marek, J. Pillet, G. Ubaghs & D. Vizcaïno (eds.):
Calymenina, Echinodermata et Hyolitha de l’Ordovicien Inferieur de la
Montagne Noire (France Meridionale). Mémoire de la Société d’Études
Scientifiques de l’Aude, 57–62.
Marek, L., 1989: The hyoliths of the Králùv Dvùr Formation (Bohemian
Ordovician). Sborník geologických ved, řada Paleontologie 30, 37–59.
Marek, L. & Yochelson, E.L., 1964: Paleozoic mollusk: Hyolithes. Science
146, 1674–1675.
Marek, L. & Yochelson, E.L., 1976: Aspects of the biology of Hyolitha
(Mollusca). Lethaia 9, 65–82.
Marek, L., Parsley, R.L. & Galle, A., 1997: Functional morphology of
hyoliths based on flume studies. Vestník Ceského geologického ústavu 72,
351–358.
Martí Mus, M. & Bergström, J., 2007: Skeletal microstructure of helens,
lateral spines of hyolithids. Palaeontology 50, 1231–1243.
Mens, K., Bergström, J. & Lendzion, K., 1987: The Cambrian system on
the East European platform (correlation chart and explanatory notes),
Valgus, Tallinn, 118 pp. [In Russian].
Mens, K., Bergström, J. & Lendzion, K., 1990: The Cambrian system on
the East European platform, correlation chart and explanatory notes.
Publication of the International Union of Geological Sciences 25, 1–73.
Meškova, N.P. & Sysoev, V.A., 1981: The discovery of traces of the digestive
system in Lower Cambrian hyoliths. In B.S. Sokolov (ed.): Problematiki
Fanerozoâ. Trudy Instituta Geologii i Geofiziki Sibirskoe Otdelenie
Akademiâ Nauk 481, 82–85 [In Russian].
Milthers, V., 1930: Bornholms geologi. Danmarks geologiske Undersøgelse,
5 Række 1, 1–140.
Missarževskij, V.V., 1969: Descriptions of hyoliths, gastropods,
hyolithelminths, camenids, and forms of an obscure systematic position.
In A.Û. Rozanov, V.V. Missarževskij, N.V. Volkova, L.G. Voronova, I.N.
Krylov, B.M. Keller, I.K. Korolûk, K. Lendzion, R. Mihnâk, N.G. Pyhova
& A.D. Sidorov: Tommotskij ârus i problema nižnej granicy kembriâ.
Akademia Nauk SSSR Trudy Geologičeskogo Instituta Akademii Nauk
SSSR, 206, 105–175. [In Russian, English translation, 1981. Paleontologic
part, 112–160. In Raben, M.E. (ed.): The Tommotian Stage and the
Cambrian Lower Boundary Problem. Amerind, New Delhi, 359 pp.].
Moberg, J.C., 1892: Om en nyupptäckt fauna i block av kambrisk sandsten,
insamlade av Dr. N.O. Holst. Geologiska Föreningens Stockholm
Förhandlingar 14, 103–120.
Moczydłowska, M. & Vidal, G., 1992: Phytoplankton from the Lower
Cambrian Læså formation on Bornholm, Denmark: biostratigraphy
and palaeoenvironmental constraints. Geological Magazine 129, 17–40.
Moysiuk, J., Smith, M.R. & Caron, J.-B., 2017: Hyoliths are Palaeozoic
lophophorates. Nature 541, 394–397. doi:https://doi.org/10.1038/
nature20804.
Nielsen, A.T. & Schovsbo, N.H., 2007: Cambrian to basal Ordovician
lithostratigraphy in Southern Scandinavia. Bulletin of the Geological
Society of Denmark 53, 47–92.
Nielsen, A.T. & Schovsbo, N.H., 2011: The Lower Cambrian of Scandinavia:
depositional environment, sequence stratigraphy and palaeogeography.
Earth Science Reviews 107, 207–310.

Pingel, C., 1828: Om overgangsformationen paa Bornholm. Tidsskrift for
Naturvidenskaberne 5, 285–297.
Poulsen, C., 1967: Fossils from the Lower Cambrian of Bornholm. Kongelige
Danske Videnskabernes Selskab, Matematisk-Fysiske Meddelelser 36,
1–46.
Poulsen, C. & Poulsen, V., 1975: Neksø sandsten og lag fra Jordens
oldtid. In A. Nørrevang & T.J. Meyer (eds.): Danmarks Natur, Bd. 1,
Landskabernes opståen, 49–82. Politikens Forlag, København.
Qian, Y. & Bengtson, S., 1989: Palaeontology and biostratigraphy of the
Early Cambrian Meishucunian Stage in Yunnan Province, South China.
Fossils and Strata 24, 1–156.
Rasmussen, H.Wienberg 1966: Danmarks geologi, 174. Gjellerup,
København.
Rasmussen, H.W., 1969: Palæontologi, fossile invertebrater, 420. København,
Munksgaard.
Rosenkrantz, A., 1938: Geologisk fører for en tre-dages ekskursion til
Bornholm. Bagges kgl, 16. Hofbogtrykkeri, København.
Runnegar, B., Pojeta, J., Morris, N.J., Taylor, J.D., Taylor, M.E. & McClung,
G., 1975: Biology of the hyolitha. Lethaia 8, 181–191.
Schiønning, E. & Wagner, P., 1967: Geologi, 120. Schultz, København.
Sinclair, G.W., 1946: Notes on the nomenclature of Hyolithes. Journal of
Paleontology 20, 72–85.
Sun, H., Babcock, L.E., Peng, J. & Zhao, Y., 2016: Three-dimensionally
preserved digestive systems of two Cambrian hyolithides (Hyolitha).
Bulletin of Geosciences 91, 51–56.
GFF   37
Sysoev, V.A., 1958: The superorder Hyolithoidea. In N.P. Luppov & V.V.
Druŝic (eds.): Principles of Palaeontology, Mollusca-Cephalopoda.
Akademiâ Nauk SSSR, 184–190. [In Russian].
Sysoev, V.A., 1962: Cambrian hyolithids from the northern slope of the
Aldan Shield. Âkutskij Filial Sibirskogo Otdeleniâ, Akademii Nauk SSSR,
66 pp. [In Russian].
Sysoev, V.A., 1973: Some features of gas chamber growth in early Cambrian
hyolithids. Paleontologičheskij Žhurnal 3, 53–57. [In Russian].
Sysoev, V.A., 1976: Terminology and methods for the investigation of
hyolithids. Paleontologičheskij Žurnal 4, 61–76. [In Russian].
Sysoev, V.A., 1984: Morphology and systematic position of the hyoliths.
Paleontologičheskij Žurnal 2, 1–10. [In Russian].
Thoral, M., 1935: Contribution à l’etude Paléontologique de l’Ordovicien
inférieur de la Montagne Noire et révision sommaire de la faune
cambrienne de la Montagne Noire, 363. Charité, Montpellier.
Valent, M., Fatka, O., Szabad, M., Micka, V. & Marek, L., 2012: Two new
orthothecids from the Cambrian of the Barrandian area (Hyolitha,
Skryje-Týřovice Basin, Czech Republic). Bulletin of Geosciences 87,
241–248.
Yochelson, E.L., 1961: The operculum and mode of life of Hyolithes.
Journal of Paleontology 35, 152–161.
Yu, W., 1974: Cambrian hyoliths. In Handbook of the Stratigraphy and
Paleontology of Southwest China, 111–113. Science Press, Beijing.
Zhao, Y. & Bengtson, S., 1999: Embryonic and post-embryonic development
of the Early Cambrian cnidarian Olivooides. Lethaia 32, 181–196.