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A revision of the Ordovician cephalopod Bactrites sandbergeri Barrande:

Systematic position and palaeobiogeography of Bactroceras

Article in Geobios · April 2015

DOI: 10.1016/j.geobios.2015.03.002

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Original article

A revision of the Ordovician cephalopod Bactrites sandbergeri

Barrande: Systematic position and palaeobiogeography of Bactroceras§
Martina Aubrechtová *
Institute of Geology and Palaeontology, Faculty of Science, Charles University in Prague, Albertov 6, 12843 Praha 2, Czech Republic

A R T I C L E I N F O A B S T R A C T

Article history: Over one hundred specimens assigned to three species of the cephalopod genus Bactroceras
Received 8 July 2014 (= Eobactrites) from the Ordovician of the Prague Basin (Czech Republic) have been studied in detail.
Accepted 26 March 2015 Preservation of the material allows study of external features including the embryonic shell, as well as
Available online 11 April 2015
some internal structures of the shell. Muscle scars, described for the first time in Bactroceras, show a
simple, unpaired and low, dorsally-situated lobe (‘‘orthoceridan’’ type of muscle scars). The type species
Keywords: of the genus, Bactroceras avus, is synonymized with Bactroceras sandbergeri based on strong similarities in
Orthocerida
the outer morphology of the shell and the position and internal structure of the siphuncle. A new species
Morphology
Embryonic shell
of Bactroceras from Bolivia is erected: Bactroceras boliviensis. Orthoceras interpolatum Barrande, 1870,
Palaeobiogeography from the Upper Ordovician of Bohemia, is assigned to Bactroceras; it is one of the stratigraphically
Prague Basin youngest species of the genus. The assignment of Orthoceras gossei Etheridge to Bactroceras is rejected.
Ordovician Convergence with bactritids and the classification of Bactroceras as the oldest known member of the order
Orthocerida is discussed. The exact stratigraphic occurrences of Bactroceras are refined, and the
palaeobiogeographic significance of the genus is evaluated. It suggests origin of the genus in high-latitude
margins of Gondwana, and its expansion to low-latitude regions during the Ordovician as part of the early
radiation of pelagic cephalopods.
ß 2015 Elsevier Masson SAS. All rights reserved.

1. Introduction represented the basal line of bactritids. Based on Bactroceras

The cephalopod genus Bactroceras Holm, 1898 is known from
the Ordovician strata of Australia, Bohemia, Bolivia, China, France,
Germany, Indonesia, Iran, Morocco, Nevada, Norway, Svalbard,
Sweden, Turkey, and Wales (Moore, 1964; Evans, 2005; Kröger and
Evans, 2011; Fig. 1). Bactroceras is characteristic in possessing a
slender, straight shell with a low angle of expansion, relatively
deep phragmocone chambers, and a narrow, ventral siphuncle. The
surface of the shell is only delicately sculptured by dense, faint
growth lines. The embryonic shell is of medium size (4 mm in
height and 2 mm in diameter) and sub-spherical in shape
(Fig. 2(B)); a cicatrix has not been observed.
The rather unique morphological features of Bactroceras led in
the past to discussions of its systematic position and evolutionary
significance. The combination of a straight shell and a narrow
marginal siphuncle in Bactrites sandbergeri Barrande, 1867 led
Schindewolf (1932, 1933) to hypothesize that the species
§
Corresponding editor: Bertrand Lefebvre.
* Corresponding author.
E mail address: aubrechm@gmail.com.
sandbergeri, he established a monotypic genus Eobactrites Schin-
dewolf, 1932. The bactritid affinity of Eobactrites was also accepted
by Erben in Moore (1964), and more recently by Holland (2003)
and Shevyrev (2006a,b). However, Flower and Kummel (1950) and
Sweet (1958) challenged the assignment of the genus Eobactrites to
the bactritids. Based on the study of Norwegian specimens, Sweet
(1958) argued that Eobactrites should rather be placed in the
nautiloid family Baltoceratidae. Sweet (1958) also implied close
affinity of Eobactrites to two additional genera of this family:
Baltoceras (= Cochlioceras) and Bactroceras. Later, other authors
(e.g., Balashov in Ruzhencev, 1962; Flower, 1964; Furnish and
Glenister in Moore, 1964; Dzik, 1981, 1984) concurred with this
opinion. Dzik (1981, 1984) and Evans (2005) synonymized
Eobactrites with Bactroceras, and placed both in the family
Baltoceratidae Kobayashi, 1935. Originally, the family was defined
as a member of the order Endocerida by Kobayashi (1935); then
the group was regarded as belonging to the order Ellesmerocer-
atida (Flower in Flower and Kummel, 1950). Mutvei (2002a,b) and
Kröger and Mutvei (2005) classified the family Baltoceratidae
within the order Orthocerida Kuhn, 1940. Especially within the last
two decades, published studies implied that Bactroceras, together

http://dx.doi.org/10.1016/j.geobios.2015.03.002
0016-6995/ß 2015 Elsevier Masson SAS. All rights reserved.
194 M. Aubrechtová / Geobios 48 (2015) 193–211

Fig. 1. Summary of known stratigraphical and geographical occurences of species of the genus Bactroceras. N. Gond.: Northern Gondwana; Av.: Avalonia; E. Gond.: Eastern
Gondwana; Laur.: Laurentia; H.: Hirnantian. Chronostratigraphical scale modified after Fatka and Mergl (2009).

with the genus Annbactroceras, are the earliest known orthocerid realms. Soon, orthocerids became highly diversified and geograph-
cephalopods (Kröger and Evans, 2011). This group includes pelagic ically widespread (Frey, 1995; Kröger and Mutvei, 2005; Kröger
cephalopods that, owing to their morphology, were able to expand et al., 2007; Kröger and Evans, 2011; Evans et al., 2013; Kröger,
from their original, demersal niches to pelagic and deep-water 2013). Consequently, understanding the origin of Bactroceras, its
 M. Aubrechtová / Geobios 48 (2015) 193–211 195

Fig. 2. Bactroceras sandbergeri; Bohemia, Czech Republic; Middle Ordovician, Darriwilian, Oretanian regional stage, Šárka Formation. A. Specimen NM L 6584, Osek u Rokycan,
lectotype. B. Specimen NM L 10331, Osek u Rokycan, juvenile phragmocone and the protoconch. C, D. Specimens MBHR 1981, Osek u Rokycan. E. Specimen CW13, Osek u
Rokycan, preserved growth lines with hyponomic sinus, originally collected and provided by V. Kozák. F. Specimen NM L 41022, Osek u Rokycan. G. Specimen NM L 40993,
Šárka-pole, living chamber with preserved muscle-scars. H. Specimen NM L 21006, Osek u Rokycan, living chamber with preserved sculpture showing hyponomic sinus.
I. Specimen NM L 41309, Zbiroh-Pětidomky, phragmocone with preserved sculpture showing hyponomic sinus. J. Specimen NM L 40994, Šárka-cihelna. K. Specimen NM L
6579, Šárka-cihelna. Scale bars: 5 mm (A, C–I, K), 1 mm (B, J).
196
evolution, and the divergence of allied lines are crucial for
elucidating the still poorly-known early radiation of orthocerid
cephalopods.
Another interesting aspect of Bactroceras is the convergence of
shell form with bactritids. Following the Ordovician radiation,
orthocerids became the most abundant cephalopod group in the
Silurian, and particularly in the Early Devonian (Kröger and Zhang,
2009), when the bactritids, the stem-group for both coleoids and
ammonoids, diverged from them (Schindewolf, 1932; Erben in
Moore, 1964; De Baets et al., 2012a, 2012b; Korn and Klug, 2003;
Kröger and Mapes, 2007; Kröger and Zhang, 2009; Kröger et al.,
2009, Kröger, 2013).
Bactroceras from the Ordovician strata of the Prague Basin,
Bohemia (Figs. 3, 4) is studied in detail in this paper. B. sandbergeri
(Barrande, 1867) occurs in siliceous nodules of the Šárka Formation
(Middle Ordovician [lower to middle Darriwilian] of the Prague
Basin, Czech Republic; Fig. 4). Barrande (1867) had a relatively small
collection at his disposal; subsequently, more than one hundred
well-preserved specimens have been collected during the 20th
century by private collectors and form the basis for the present
study. Kraft and Kraft (1994) and Marek (1999) noted the occurrence
of Bactroceras cf. sandbergeri in shales of the Klabava Formation
(Floian to Dapingian; Fig. 4) – a formation preceding the Šárka
Formation. Apart from the Prague Basin, fragments of B. sandbergeri
have also been reported from strata of a similar age in the Oslo
region, Norway (Sweet, 1958), and in Wales, United Kingdom
(Evans, 2005) (Figs. 1, 5). In addition to B. sandbergeri, Barrande
described Orthoceras interpolatum Barrande, 1870 from the Late
Ordovician Králův Dvůr Formation (upper Katian to lowermost
Hirnantian; Fig. 4) of Bohemia. Herein, this species is assigned to
Bactroceras. It is notable that Bactroceras occurs in the Prague Basin
in the Klabava, Šárka and Králův Dvůr formations, which contain
faunas with Baltic affinities (Havlı́ček et al., 1994; Fatka and Mergl,
2009). Examination of the distribution pattern and dispersion of
Bactroceras may thus help in reconstructions of migration seaways
in the Ordovician (Havlı́ček et al., 1994; Fatka and Mergl, 2009).
Fig. 3. Map of the Prague Basin showing distribution of Ordovician rocks and localities from which Bactroceras is known. 1. Klabava (Starý hrad), Klabava Formation;
2. Rokycany (Drahouš), Šárka Formation; 3. Osek near Rokycany, Šárka Formation; 4. Dı́ly u Rokycan, Šárka Formation; 5. Mýto near Rokycany, Šárka Formation; 6. Cekov,
Šárka Formation; 7. Kařez, Šárka Formation; 8. Kařı́zek, Šárka Formation; 9. Zbiroh (Pětidomky), Šárka Formation; 10. Újezd, Šárka Formation; 11. Lejškov near Málkov, Králův
Dvůr Formation; 12. Králův Dvůr, Králův Dvůr Formation; 13. Praha (Šárka), Šárka Formation; 14. Popovice, Šárka Formation. Map modified after Manda (2008).
M. Aubrechtová / Geobios 48 (2015) 193–211
2. Material and methods
The material studied consists of over one hundred specimens.
The stratigraphically oldest material is from the Klabava
Formation (Floian to Dapingian; Fig. 4). This material includes
flattened fragments of phragmocones preserved in green-grey
shales. The bulk of the material comes from the overlying Šárka
Formation (lower to middle Darriwilian; Fig. 4). This material
consists of external and internal moulds of body-chambers and
phragmocones, mostly preserved in siliceous nodules. Specimens
are rarely found in the shales. Specimens from the Šárka
Formation are usually not deformed. The shell wall is in many
specimens at least partly preserved and coated with oxides and
hydroxides of iron. Original internal structures of the shell
(septal necks, connecting rings) are fully dissolved. The
stratigraphically youngest specimens assigned to the genus
were recorded from the Králův Dvůr Formation (upper Katian to
lowermost Hirnantian; Fig. 4). Here, a living chamber and a part
of a phragmocone are preserved in carbonate nodules and dark-
grey siltstones.
The material was described and measured in detail using an
Olympus SZX-12 stereoscopic microscope. About 30 specimens
were coated with ammonium chloride and photographed, using an
Olympus DP72 camera attached to the microscope. Latex casts
have been made of several of the best-preserved specimens in
order to obtain details of shell sculpture, muscle scars, shell
damages and siphuncular structure. Images of these specimens
were processed using Adobe Illustrator CS 11.0.0., CorelDRAW X4
and X6, and Adobe Photoshop CS6.
The morphological terminology follows Teichert (in Moore,
1964). Institutional abbreviations and prefixes: NM L (National
Museum, Prague); CGS CW (Czech Geological Survey, Prague); UK
CHMHZ (Charles University, Prague, Faculty of Science); UK UGP
(Charles University, Prague, Faculty of Science); MBHR (Dr.
Bohuslav Horák Museum, Rokycany); NRS MO (Naturhistoriska
Riksmuseet, Stockholm, Sweden).
 M. Aubrechtová / Geobios 48 (2015) 193–211 197

Diagnosis (comp. Kröger and Isakar, 2006): Orthoconic to

cyrtoconic, smooth to elaborately ornamented shells. Siphuncle
typically narrow, central or subcentral, but in early evolutionary
forms may be wide and marginal in position. Siphuncle tubular or
slightly expanded between chambers. Septal necks achoanitic,
suborthochoanitic, orthochoanitic, or hemichoanitic. Apex small to
medium-sized, spherical or sub-spherical, bowl-shaped or blunt,
invariably without cicatrix, slightly constricted or unconstricted.
Cameral deposits rather well developed to strongly suppressed.
Endosiphuncular deposits not present or strongly suppressed,
developed in early evolutionary forms. Muscle scars with unpaired
or paired, low, dorsal, anterior lobe or lobes.
Remarks: The importance of internal structures of the shell for
classification at generic or higher taxonomic levels was stressed by
Hyatt (1883-1884). His concepts altered the morphological
classification of orthocerids, which had previously placed emphasis
on the gross shell form. However, Hyatt’s concept was neglected for
a long time, especially by European palaeontologists (e.g.,
Troedsson, 1926). Hyatt’s views were further elaborated by Flower
(1962), who studied the internal morphology of these taxa in more
detail. Ruzhencev (1962) and Sweet in Moore (1964) summarized
the knowledge on cephalopod morphology and views on their
systematics. The latter authors assigned the majority of straight-
shelled cephalopods to the order Orthocerida, which comprised two
superfamilies: Orthocerataceae and Pseudorthocerataceae. How-
ever, in the recent decades, other, previously neglected characters
have been considered when discussing the systematic position of
Fig. 4. Stratigraphical scheme of the Ordovician of the Prague Basin, Bohemia. the Orthocerida. As a consequence, different classifications of
Stratigraphical levels at which the genus Bactroceras has been recorded are marked orthocerid cephalopods were offered, and the group in the concept
in grey color. Per.: Perunica. Modified after Ernst et al. (2014).
of Ruzhencev (1962) and Sweet in Moore (1964) was shown to be
polyphyletic (e.g., Flower, 1964; Zhuravleva, 1994; Evans, 2005).
3. Systematic palaeontology The new concepts demonstrated that crucial characters for the
systematics of orthocerids include not only the detailed structure of
Class CEPHALOPODA Cuvier, 1798 the siphuncle, but also the character of the embryonic shell and type
Subclass ORTHOCERATOIDEA Kuhn, 1940 of muscle scars. Orthocerid cephalopods that are characterized by
Order Orthocerida Kuhn, 1940 short septal necks, thin connecting rings and in many lineages
Included Families: see Evans (2005), Kröger and Isakar (2006), suppressed endosiphuncular and cameral deposits (Barskov, 1963),
and Kröger et al. (2007). also display single-layered septal necks (Zakharov, 1996), connect-

Fig. 5. Palaeogeographic distribution of the genus Bactroceras during the Middle Ordovician. Map modified after Cocks and Torsvik (2006).
198 M. Aubrechtová / Geobios 48 (2015) 193–211
ing rings with pores (Mutvei, 2002a,b), muscle scars with a low,
dorsal lobe or lobes (Mutvei, 2002a,b) and, most importantly, a
small to medium-sized subspherical or spherical embryonic shell
without a cicatrix (e.g., Balashov, 1957; Ristedt, 1968; Kolebaba,
1973; Engeser, 1996; Kröger, 2006). Thus, it can be seen that the
mentioned features correlate very well with each other, and differ
significantly from the corresponding structures described in other
groups of straight-shelled cephalopods (Engeser, 1996). Moreover,
these features are also present amongst the oldest known straight-
shelled cephalopods reported from various Early and Middle
Ordovician rocks, namely Bactroceras, studied herein. The system-
atic position of the genus was uncertain until its embryonic shell
was described by Dzik (1981, 1984) and Evans (2005) for two
species of the genus: B. sandbergeri and B. angustisiphonatum. The
embryonic shell of B. sandbergeri was first illustrated by Barrande
(1870) – who, however, referred it to Tretoceras parvulum Barrande).
The protoconch (Fig. 2(B)) of the species is medium-sized (4 mm in
height and 2 mm in diameter), subspherical and lacks a cicatrix. In B.
angustisiphonatum, Evans (2005) described and illustrated the
embryonic shell as possessing a small (1.25 mm in height, 1.1 mm in
diameter), spherical protoconch. Mutvei (2002a,b) described
‘‘orthoceridan’’ (i.e., porous) connecting rings in Bactroceras. In
addition, ‘‘orthoceridan’’ muscle scars imprints (according to the
terminology used by Mutvei, 2002a,b) have been discovered in
B. sandbergeri (Fig. 2(G)).
The results of the present study are in accordance with current
concept of the order Orthocerida as defined by Kröger and Isakar
(2006). In their view, Orthocerida now constitutes a monophyletic
taxon and of the characters listed, the form of the embryonic shell
is the most important for elucidating the systematics of the order
(Kröger and Mapes, 2004, 2007; Evans, 2005; Turek, 2010; Turek
and Manda, 2012).
Family BALTOCERATIDAE Kobayashi, 1935
Diagnosis (After Kröger et al., 2007): Orthocones with nearly
straight sutures and septal spacing with orthoceridan aspect (= high
phragmocone chambers, two or three chambers correspond to shell
diameter). Siphuncle marginal, or located between center and
margin in some advanced forms. Siphuncular tube generally wide,
in some forms narrow; tubular or slightly expanded between septa;
no diaphragms present. Connecting ring thin compared with
ellesmeroceridans and Sactorthoceratidae. Septal necks subortho-
choanitic or orthochoanitic. Apex spheroidal, with constriction.
Endosiphuncular deposits consist of tubular calcareous rods
typically depressed circular in section. Cameral deposits typically
mural, episeptal, restricted to adapical half of phragmocone.
Remarks: The Early Ordovician is a crucial period for
understanding the early evolution of orthocerids and their
descendants, as the oldest members of this group first appeared
at that time. These are now included in the family Baltoceratidae.
Many other broadly coeval cephalopods possessing slender,
orthoconic shells have been described (e.g., Evans, 2005; Kröger
et al., 2007); however, the systematic position of these is often
unclear due to poor preservation and lack of some of the diagnostic
characters. These forms are mostly assigned to orders Dissidocer-
ida and Ellesmerocerida (Evans, 2005; Kröger et al., 2007; Kröger
and Evans, 2011).
The family Baltoceratidae was defined by Kobayashi (1935) as a
member of the order Endocerida. Flower in Flower and Kummel
(1950) assigned the family to the Ellesmerocerida. This approach was
followed widely in the next decades (e.g., Flower, 1964; Furnish
and Glenister in Moore, 1964; Dzik, 1981, 1984). However, as a
consequence of the recent revision of the Ellesmerocerida by Kröger
and Mutvei (2005), the family Baltoceratidae was placed in the order
Orthocerida. This classification is supported here, as members of the
family, including Bactroceras, bear orthoceridan diagnostic features.
A closer look at the various genera classified within the family
Baltoceratidae showed that the group was actually polyphyletic
(Zhuravleva, 1994; Evans, 2005). Recently, in order to resolve this
situation, Kröger et al. (2007) studied Dapingian and Darriwilian
cephalopod assemblages of the Argentine Precordillera, emending
the family Baltoceratidae and re-assigning some of the genera that
previously belonged to this family. The morphology of the genus
Bactroceras from the Prague Basin corresponds to the generic
diagnosis of Kröger et al. (2007); it is therefore regarded as a
member of Baltoceratidae. In Bactroceras, the siphuncle is marginal
or almost marginal, the shell possesses high phragmocone
chambers, no diaphragms are present, siphuncular segments are
convex, and septal necks are short. However, other genera that are
currently also assigned to that group, namely the type genus
Cochlioceras, exhibit a different morphology, as they have a rather
wide siphuncle, relatively short phragmocone chambers, and often
possess endosiphuncular deposits. Thus, several genera which are
presently assigned to the Baltoceratidae remain in need of future
revision in order to determine whether the family is indeed
monophyletic or needs further redefinition.
Genus Bactroceras Holm, 1898
Type species: Bactroceras avus Holm, 1898; Middle Ordovician,
upper Darriwilian, Seby Limestone Formation; Öland, Sweden.
Included species: Bactroceras angustisiphonatum (Rüdiger,
1889); Grey Lituites Limestone (Lasnamägian Folkeslunda Lime-
stone), Middle Ordovician, upper Darriwilian; Mecklenburg,
Germany. Bactroceras boliviensis nov. sp.; Pircancha Formation,
Chaupiuno, Lower Ordovician, upper Floian; Tarija, Bolivia.
Bactroceras huanghuaense Xu and Lai, 1987; Middle Ordovician;
Yangtze Gorge area, China. Bactroceras interpolatum (Barrande,
1870) nov. comb.; Upper Ordovician, upper Katian; Prague Basin,
Czech Republic. Bactroceras mourguesi (Thoral, 1935); Lower
Ordovician, upper Tremadocian/lower Floian; Montagne Noire,
France. Bactroceras sandbergeri (Barrande, 1867); Middle Ordovi-
cian, Darriwilian, Oretanian, Šárka Formation; Prague Basin, Czech
Republic. Bactroceras cf. sandbergeri (Barrande, 1867); Middle
Ordovician, upper Dapingian, Klabava Formation; Prague Basin,
Czech Republic. Bactroceras sp. (Kröger and Lefebvre, 2012); Lower
Ordovician, upper Tremadocian, Lower Fezouata Formation, H.
copiosus Biozone; Anti-Atlas, Zagora area, Morocco. ?Bactroceras sp.
(Dean and Monod, 1970); Lower Ordovician, upper Tremadocian-
lower Floian, Seydisehir Shales; Taurus Mountains, Turkey.
?Bactroceras sp. (Evans et al., 2013); Lower Ordovician, upper
Tremadocian; Eastern Alborz, Iran. Bactroceras subventrum Lai,
1987; Upper Ordovician, lower Sandbian; South China. Bactroceras
xianlingbuense Zou, 1987), Middle Ordovician, middle Darriwilian,
Kuniutan Formation; South China. Bactroceras zhejiangense Zou,
1987; Middle Ordovician, middle Darriwilian, Kuniutan Forma-
tion; South China.
Diagnosis (comp. Kröger and Evans, 2011): Slender, straight or
slightly curved shell with sub-circular or circular cross-section,
apical angle low, ranges between 58 and 108. Surface sculpture
with straight, transverse growth lines or annulations, and faint
striae. Hyponomic sinus broad and shallow. Phragmocone
chambers rather deep, concavity at least 30%. Sutures straight
and transverse. Distance between two sutures between 1/2 and 1/4
that of the phragmocone diameter. Siphuncle marginal or slightly
removed from shell margin; diameter of the siphuncle between 1/6
and 1/20 that of the phragmocone. Septal necks orthochoanitic or
hemichoanitic; siphuncular segments tubular or slightly expanded
within phragmocone chambers; connecting rings thin and
homogeneous. Endosiphuncular and cameral deposits not present.
Remarks: Bactroceras was originally described by Holm (1898)
based on B. avus Holm, 1898 (Fig. 6(E–H)) from the Darriwilian
rocks of Öland, Sweden, which is regarded here as a junior
 M. Aubrechtová / Geobios 48 (2015) 193–211 199

Fig. 6. A–D, G. Bactroceras boliviensis nov. sp.; Chaupiuno, Tarija, Bolivia; Lower Ordovician, Floian, Pircancha Formation. Specimen UGP-O-0001 (holotype), phragmocone (C,
D) showing shell sculpture (A), section showing internal structure of phragmocone (B), and detail of the siphuncle (G); originally collected and provided by B. Weber. E, F, H.
Bactroceras sandbergeri (= Bactroceras avus); Sweden; Middle Ordovician, Darriwilian, Seby Limestone Formation. E, F: Specimen NRS MO 154302, Öland, phragmocone with
preserved structure of the siphuncle; H: Specimen NRS MO 154302, Öland, thin section. Scale bars: 5 mm.

synonym of B. sandbergeri (see below). The species differs from B. angustisiphonatum (see Rüdiger, 1889; Zou, 1987; Evans,
other species mainly in the presence of a very narrow siphuncle, 2005) differs remarkably from B. sandbergeri in having wider septal
which is nearly in contact with the shell wall and the segments of spacing and larger diameter of the siphuncle (1/6 to 1/4 that of the
which are slightly expanded (Fig. 6(F, H)). phragmocone; Figs. 7, 8). Also, the siphuncle is submarginal and
200 M. Aubrechtová / Geobios 48 (2015) 193–211
Fig. 7. Bivariate diagram showing variations in siphuncle proportional diameter in
species of the genus Bactroceras.
rather more tubular (Fig. 8). Concavity of septa is only 1/3 that of
the following phragmocone chamber (Fig. 8). Transverse adorally,
imbricated striae are present on the surface of the shell. In addition,
the embryonic shell described by Evans (2005) is much smaller in
diameter and much more spherical in shape than that of B.
sandbergeri (Dzik, 1981, 1984; Evans, 2005). Morphologically
similar to B. angustisiphonatum is B. interpolatum nov. comb.
(Fig. 9(A, B)), based on Barrande’s (1870) O. interpolatum. The only
difference between the morphology of the two species is a
narrower siphuncle in B. interpolatum nov. comb. (Figs. 7, 8).
Besides B. angustisiphonatum, Zou (1987) described two other
species from the Darriwilian Kuniutan Formation of South China. B.
xianlingbuense is similar to B. sandbergeri in the presence of a very
narrow marginal siphuncle with slightly expanded connecting
rings (Fig. 8). B. zhejiangense is rather similar in morphology to B.
angustisiphonatum, as its shell possesses a tubular siphuncle;
however, it is marginal in position (Fig. 8).
So far, B. mourguesi from the Lower Ordovician of the Montagne
Noire, France, is the oldest known species assigned to the genus
(Kröger and Evans, 2011). Its shell differs significantly from the
shell of other species in being slightly curved in apical portions.
Also, the surface of the shell displays faint, transverse striae or
annulations, and the siphuncle of the species is narrower in
diameter than in the previously discussed species (Fig. 8).
Fig. 8. Table showing main morphological features and their distribution among some of the species of the genus Bactroceras. Features in bold are considered as derived and
are indicated with black circles (two black circles indicate extreme values of the character); features in normal letters are considered as primitive and are indicated with
empty circles.
Marek et al. (2000) mentioned specimens from the upper Floian
rocks in Chaupiuno, Bolivia, and classified them as B. aff. avus. The
general morphology of the specimens, especially the wide septal
spacing and a rather narrow, marginal siphuncle (Fig. 8) are in
agreement with the generic diagnosis. However, the siphuncle
shown by these specimens is more marginal, more tubular and
wider than that in B. avus (Fig. 8). Also, the specimens are too large
in proportions to be assigned to B. avus or any other species of the
genus. Thus, they are herein described as a separate, new species B.
boliviensis (Fig. 6(A–D, G)).
Specimens belonging to Bactroceras were described from
Morocco (Kröger and Lefebvre, 2012) and Iran (Evans et al.,
2013). These authors reported that the studied specimens show
similarities with B. angustisiphonatum and B. avus, respectively, but
because of the lack of diagnostic characters, no species-level
determination was possible. Based on the descriptions and
photographs provided by Kröger and Lefebvre (2012), the
specimen from Morocco probably belongs to B. angustisiphonatum,
as its phragmocone chambers are too long and siphuncle too wide
for any other species. However, as neither the surficial ornamen-
tation nor the expansion rate are known, exact determination at
the species level remains impossible (Kröger and Lefebvre, 2012).
The Iranian specimen assigned to the genus Bactroceras by Evans
et al. (2013) is a small portion of a phragmocone, showing three
phragmocone chambers and a marginal siphuncle. However, the
phragmocone chambers are very short for Bactroceras; thus, the
present author considers the generic assignment of this specimen
to Bactroceras questionable.
Etheridge (1893) described Orthoceras gossei from the Darri-
wilian of Australia. Teichert and Glenister (1952) assigned the
species to Bactroceras. The type specimen illustrated by Etheridge
(1893) shows a rather narrow and marginal siphuncle; however, it
possesses very short phragmocone chambers, and thus clearly does
not belong to Bactroceras.
B. huanghuaense Xu and Lai, 1987 from the Middle Ordovician of
the Yangtze Gorge area, China, differs from all other species in
having an extremely narrow, tubular siphuncle (1/26 of the
phragmocone diameter; Fig. 8), which is significantly shifted from
the shell wall (distance from shell is 4 mm).
As stated by Evans (2005), distinguishing between individual
species of Bactroceras is often difficult, due to poor preservation
and unpreserved species-specific diagnostic characters. This
comment mainly applies to the species B. subventrum (Lai,
1987), B. interpolatum (Barrande, 1870) nov. comb., B. xianling-
buense (Zou, 1987), and Bactroceras zhejiangense (Zou, 1987).
Although these species clearly belong to Bactroceras, their species
status remains uncertain.
 M. Aubrechtová / Geobios 48 (2015) 193–211 201

Fig. 9. A, B. Bactroceras interpolatum nov. comb.; Bohemia, Czech Republic; Upper Ordovician, upper Katian, Králův Dvůr Formation. A: Specimen NM L 20994, Králův Dvůr; B:
Specimen NM L 13764, Lejškov u Málkova. C–K. Bactroceras sandbergeri; Bohemia, Czech Republic; Middle Ordovician, Darriwilian, Oretanian regional stage, Šárka Formation.
C: Specimen CGS 451, PP 546, Osek u Rokycan, sublethal damages of the shell; D: Specimen NM L 41001, Kařı́zek; E: Specimen NM L 40999, Praha-Šárka; F-H: Specimen NM L
41002, Cekov, phragmocone (F), latex cast (G), detail (H); I: Specimen NM L 6577, Osek u Rokycan; J: Specimen NM L 40996, Praha-Šárka, sublethal damages of the shell; K:
Specimen NM L 21005, Osek u Rokycan, wrinkled layer. Scale bars: 1 mm (A, K), 5 mm (B–J).
202 M. Aubrechtová / Geobios 48 (2015) 193–211
Bactroceras sandbergeri (Barrande, 1867)
Figs. 2, 6(E, F, H), 9(C–K)
1867. Bactr. Sandbergeri Barr. - Barrande, pp. 49-50.
1868. Bactr. Sandbergeri (Barr.) - Barrande, pl. 245, figs. 9-21.
1868. Tretoceras parvulum Barr. - Barrande, pl. 247, figs. 26-28.
1870. Bactrites Sandbergeri (Barr.) - Barrande, pl. 413, figs. 10-14.
1870. Orthoceras naufragum Barr. - Barrande, pl. 415, figs. 6-8,
9-10.
1874. Tretoceras parvulum (Barr.) - Barrande, p. 801.
1889. Endoceras nov. sp. - Rüdiger, p. 36, pl. 1, fig. 5a-b.
1898. Bactrites sandbergeri (Barr.) - Holm, pp. 355–358.
1898. Bactroceras avus Holm - Holm, pp. 358–359, pl. 1, figs. 1-7.
1932. Eobactrites sandbergeri (Barr.) - Schindewolf, pp. 173-174.
1933. Eobactrites sandbergeri (Barr.) - Schindewolf, pp. 67-73, pl.
4, fig. 9a, b.
1934. Eobactrites sandbergeri (Barr.) - Schindewolf, pp. 270-280
1950. Eobactrites sandbergeri - Flower and Kummel, p. 608.
1958. Eobactrites sandbergeri (Barrande) - Sweet, pp. 28-30, pl.
2, figs. 1, 3, 5.
1960. Eobactrites sandbergeri (Barrande) - Špinar, p. 456.
1962. Eobactrites sandbergeri (Barrande) - Ruzhencev, p. 75,
fig. 38.
1964. Eobactrites - Furnish and Glenister in Moore, K132.
1964. Eobactrites Schindewolf (= Bactrites sandbergeri Barrande,
1867) - Erben in Moore, K495, K501, fig. 358 (2a, b).
1964. Eobactrites sandbergeri (Barrande) - Flower, pp. 107-108,
fig. 37.
1965. Eobactrites sandbergeri (Barrande) - Špinar, p. 457, figs.
8-158.
1981. Bactroceras sandbergeri (Barrande) - Dzik, p. 162.
1981. Tretoceras parvulum (Barrande) - Dzik, p. 162, fig. 1.
1984. Bactroceras (Holm) (= Eobactrites Schindewolf, 1932) -
Dzik, p. 18.
1984. Tretoceras parvulum (Barrande) (= Bactrites sandbergeri
Barrande, 1867) - Dzik, p. 18.
1985. Eobactrites sandbergeri - Hewitt and Stait, pp. 230-231.
1999. Eobactrites sandbergeri (Barrande) - Marek, p. 413.
2003. Eobactrites sandbergeri (Barrande) - Holland, pp. 370-371.
2005. Tretoceras parvulum (Barrande) - Evans, pp. 30-32.
Partim. 2005. Bactroceras sandbergeri (Barrande) - Evans, pp.
31–32, pl. 4, fig. 6, non pl. 3, fig. 19, pl. 4, fig. 1.
2006b. Eobactrites sandbergeri (Barrande) - Shevyrev, p. 155.
2007. Bactroceras Holm (= Eobactrites Schindewolf, 1932) -
Kröger and Mapes, pp. 320-321, fig. 4c.
2008. Bactroceras sandbergeri (Barrande) - Manda, p. 328.
2008. Tretoceras parvulum (Barrande) - Manda, p. 328.
Lectotype: Designated herein, specimen NM L 6584, figured by
Barrande (1867: pl. 245, fig. 18), herein re-figured in Fig. 2(A).
Type locality and horizon: Osek u Rokycan, central Bohemia,
Czech Republic; Ordovician, Darriwilian, Oretanian, Šárka Forma-
tion, Corymbograptus retroflexus Biozone.
Material: 117 studied specimens (Table 1), mostly fragments of
shells preserved as internal moulds. All specimens are stratigra-
phically confined to the lower part of the Šárka Formation. Precise
stratigraphic position is mostly not possible to determine as most
of the specimens come from old collections of siliceous nodules
weathered out of the shale outcrops. Faunal associations and
previous biostratigraphic investigations at individual localities
with Bactroceras sandbergeri mostly imply the Corymbograptus
retroflexus Biozone, or alternatively, the Didymograptus clavulus
Biozone (see Lajblová and Kraft, 2014; Table 2).
Eighty-five specimens are held in NM (protoconch L 10331;
external mould L 40996; body-chambers L 21005, L 21006, L 6580, L
40993, L 40998, L 41007, L 41008, L 41010, L 41012, L 41014–L
41016, L 41024, L 41027; fragments of phragmocones L 6576-L6579,
L 6580-L 6585, L 20993, L 20994, L 40988-L 40992, L 40994, L 40995,
L 40997, L 40999, L 41000-L 41006, L 41009, L 41011, L 41013, L
41017-L 41023, L 41025, L 41026, L 41309, L 42945-L 42953),
19 specimens were studied at MBHR (body-chamber 1901,
fragments of phragmocone 1939, 1951, 1963, 1964, 1972, 1975,
1979, 1981, 1983, 1988, 1993, 5515, 5825, 9535, 13239, 15521,
17825, 20333), 12 specimens are from CGS (internal moulds,
collectively as PP545–PP547, PP595, CW11 and CW13), and one
fragment of phragmocone (CHMHZ-OR-0001) is from UK.
Occurrence (Figs. 1, 3–5; Table 2): Prague Basin, central
Bohemia, Czech Republic; Ordovician, Darriwilian, Oretanian,
Šárka Formation; Corymbograptus retroflexus Biozone: localities
Cekov, Dı́ly u Rokycan, Kařı́zek, Mýto u Rokycan, Osek u Rokycan,
Popovice, and Zbiroh-Pětidomky; Corymbograptus retroflexus or
Didymograptus clavulus Biozone: localities Praha-Šárka, Šárka-
cihelna, Šárka-pole, Přı́lepy, and Úvaly. United Kingdom, Wales;
Ordovician, Darriwilian, Abereiddian Stage, Didymograptus artus
Biozone: locality Cefn-yr-Owen-Uchaf. Norway; Ordovician, Dar-
riwilian, upper part of the Didymograptus shales: localities Bygdøy
Sjøbad, Villa Victoria. Germany, Middle Ordovician, upper Darri-
wilian, Grey Lituites Limestone, Lasnamägian Folkeslunda Lime-
stone: locality Mecklenburg.
Measurements: See Table 1.
Diagnosis: Bactroceras with a very narrow marginal siphuncle
(diameter <10% of the phragmocone diameter). Septal necks
orthochoanitic or hemichoanitic. Connecting rings slightly ex-
panded within chambers. The embryonic shell is moderately large,
subspherical and with constriction. Surface sculpture with faint,
transverse growth lines, no annulations or striae present.
Hyponomic sinus either not developed or broad and shallow.
Apical angle ranges between 58 and 68. Concavity of septa high, up
to 1/2 that of the following phragmocone chamber.
Description: The lectotype (NM L 6584, a part of a phragmo-
cone; Fig. 2(A)) is 22.4 mm in height and 10 mm in diameter. Four
phragmocone chambers are preserved, all of them equal in height
(1/2 of the diameter of the phragmocone). The sutures are straight,
directly transverse. The siphuncle is narrow and ventral, only
slightly removed from the shell wall. No surficial structures are
preserved.
The remaining material consists of parts of phragmocones and
living chambers, ranging in diameter from 2 to 26.9 mm. The shell
is straight and circular in cross-section. The angle of expansion
ranges between 48 and 68. The sutures are straight and directly
transverse. 1.4 to 3 phragmocone chambers correspond to the
respective shell diameter; in parts of shells representing late
ontogenetic stages, 4–6 chambers correspond to the respective
shell diameter (Figs. 9(D, E), 10). The siphuncle is ventral, almost
marginal, only slightly removed from the shell wall. Cross-section
of the siphuncle is circular or very slightly laterally flattened. Its
diameter ranges between 1/10 and 1/22 of the phragmocone
diameter (Fig. 11). Septal necks are ortho- or hemichoanitic, and
closer to the shell wall than connecting rings. Connecting rings are
thin, homogeneous and slightly expanded within chambers (Figs.
2(D, J), 6(E, F, H), 9(F–H)). Ultrastructure was not observed. The
thickness of the ring wall corresponds to 1/6 of the siphuncle
diameter. The height of the most complete living chamber is
3 times the maximum shell diameter (NM L 42948). The aperture is
straight and open. Muscle scars show an unpaired lobe on the
dorsal side of the living chamber (NM L 40993; Figs. 2(G), 12). The
lobe of the annular elevation is 1/8 of the height of the living
chamber. Sculptures, where preserved (Fig. 2(E, H, I)), consist of
dense, faint growth lines (7 growth lines per mm), transverse to the
longitudinal axis of the shell. They are straight, forming a broad
and shallow lobe (hyponomic sinus NM L 21006, NM L 41309, CGS
CW13; Fig. 2(E, H, I)) on the ventral side of the shell. The lobe depth
is 1/13 of the height of the corresponding phragmocone chamber.
The protoconch (specimen NM L 10331; Fig. 2(B)) is of medium size
 M. Aubrechtová / Geobios 48 (2015) 193–211 203

Table 1
Measurement table.

Specimen Length of the Maximum shell Diameter of Diameter of

preserved part of diameter (mm) shell/height siphuncle/Diameter
the phragmocone (mm) of chamber of shell

Bactroceras sandbergeri
NM L 10331 (embryonic shell) 4 2 2 1/5
NM L 20993 63.4 16.6 3 n.a.
NM L 40988 39 12 2 n.a.
NM L 40989 17.3 9.8 3 n.a.
NM L 40990 16.2 13.9 3 1/12
NM L 40991 15 22 n.a. 1/14
NM L 40992 19 26.2 n.a. 1/12
NM L 40993 22.7 15.9 Living chamber 1/16
NM L 40994 29.2 21.2 2.7 1/21
NM L 40995 13 12 2 n.a.
NM L 40997 6.65 5.4 1.7 n.a.
NM L 40998 38.2 11.2 2.3 n.a.
NM L 40999 46.4 24.6 3 1/15
NM L 41000 24.4 13 2 1/17
NM L 41001 71.29 23.2 3.73–6 1/17
NM L 41002 57 20.3 3.2 1/14
NM L 41003 49.9 18.4 3.3 1/14
NM L 41004 24.4 20.5 3.4 1/18
NM L 41005 36 17.8 3.3 1/12
NM L 41006 28 18.9 3.5 1/22
NM L 41007 21.3 12.6 Living chamber Living chamber
NM L 41008 33 21.5 3 n.a.
NM L 41009 8 8.1 n.a. 1/14
NM L 41010 23 9.7 Living chamber Living chamber
NM L 41011 30 21 3 1/11
NM L 41012 18.1 4 Living chamber 1/20
NM L 41013 28.9 9.9 2.5 n.a.
NM L 41014 20 8.1 Living chamber Living chamber
NM L 41015 41.6 15.8 Living chamber 1/15
NM L 41016 11 15 n.a. n.a.
NM L 41017 54 11.1 1.6 1/10
NM L 41018 24.9 22 4 n.a.
NM L 41019 24 10.2 2.2 1/17
NM L 41020 15.1 8 1.7 1/1
NM L 41021 18 20 n.a. n.a.
NM L 41022 25.7 9.9 1.5–4.3 n.a.
NM L 41023 20.7 10.9 2 1/10
NM L 41024 18.7 11.9 Living chamber Living chamber
NM L 41025 33.4 8.4 2.3 1/20
NM L 41026 63.5 15.2 3.4 1/16
NM L 41027 30.9 14.3 Living chamber Living chamber
NM L 41309 45.9 19 3.5 1/22
NM L 42945 58.7 19.7 2.8 1/11
NM L 42946 24 9 1.5 1/10
NM L 42947 14.7 11.9 n.a. 1/10
NM L 42948 54.9 16.4 Living chamber 1/13
NM L 42949 41 22.4 3.5 1/18
NM L 42950 16.8 6.6 1.7 1/10
NM L 42951 17.3 4.4 1.4 1/12
NM L 42952 10.7 9.3 1.6 1/13
NM L 42953 8.9 5 2 n.a.
NM L 6576 29.2 15.2 3 1/10
NM L 6577 45.2 10.5 2 1/11
NM L 6578 50.5 13.9 2.4 n.a.
NM L 6579 29.3 26.9 3 1/13
NM L 6580 38 23.9 Living chamber Living chamber
NM L 6581 22.2 8 1.4 1/12
NM L 6582 34.3 18 2 1/15
NM L 6583 65 17 4 1/19
NM L 6584a 22.37 10 2.3 1/15
NM L 6585 39.6 22.9 4 1/20
MBHR 1951 38 22 n.a. 1/20
MBHR 1975 34 20 3 1/13
MBHR 1981 20 4 1.5 n.a.
MBHR 9535 14 9 2 1/15
MBHR 17825 35 25 3 1/20
CGS PP546 16 8 2 n.a.
UK CHMHZ-OR-0001 33 13 2 1/18

Bactroceras cf. sandbergeri

MBHR 20279 70 8 1.6 1/10
204 M. Aubrechtová / Geobios 48 (2015) 193–211
Table 1 (Continued )
Specimen Length of the
preserved part of
the phragmocone (mm)
Bactroceras interpolatum nov. comb.
NM L 13764 35.9
NM L 20994b 11.1
Bactroceras boliviensis nov. sp.
UK UGP-O-0001b 88
UK UGP-O-0004 64.5
n.a.: not available.
a
Lectotype.
b
Holotype.
(4 mm in height and 2 mm in diameter) and hemispherical in
shape, with only a slight constriction (the diameter of initial
chamber declines adorally from 2.5 to 2 mm). The second
phragmocone chamber reaches a depth 1/2 that of the phragmo-
cone diameter; the siphuncle diameter is 1/5 of the total diameter
of the phragmocone. Septal neck of the first siphuncular segment is
orthochoanitic. Neither cicatrix nor surface sculpture have been
found. The caecum is cylindrical, about 1 mm in length, and
expands slightly within the initial chamber. Height of the second
phragmocone chamber corresponds to 1/2 of the phragmocone
diameter. Cameral and endosiphuncular deposits are unknown.
Wrinkled-layer has been found at the living chamber of specimen
NM L 21005 (Fig. 9(K)). The conchal furrow (Fig. 9(F–H)) consists of
a narrow, discontinuous ridge situated mid-ventrally on the
internal mould. It is in width 1/24 (0.3 mm) of the height of the
phragmocone chamber.
Remarks: General proportions of the shell and position of the
siphuncle of Bactroceras sandbergeri from the Darriwilian of the
Prague Basin are comparable to those of Bactroceras avus. Small
differences in morphology clearly fall within the species
variability, especially when considering that only limited
material of B. avus is available so far. The remarkable morpho-
logical similarities, as well as corresponding stratigraphic
occurrence, suggest that B. avus and B. sandbergeri are actually
a single species.
Besides the Prague Basin, B. sandbergeri was also reported from
localities in Baltica and Avalonia (Figs. 1, 5). Sweet (1958)
described specimens from Darriwilian rocks at Bygdøy Sjøbad
and Villa Victoria near Oslo, Norway. All three fragments of
phragmocones show remarkable similarities with the Bohemian
material, and very probably belong to B. sandbergeri. Evans (2005)
also reported the species from the Darriwilian of Cefn-yr-Owen-
Uchaf in Wales. Other specimens, NMW2001.7G.61 (pl. 3, fig. 19)
and NMW2001.7G.61 (pl. 4, fig. 1) from the Pont-y-Fenni
Formation (Dapingian), do not belong to B. sandbergeri based on
photographs and descriptions given by Evans (2005). Although
these specimens display a narrow and marginal siphuncle, their
camerae are very low for a B. sandbergeri. On the other hand, the
third specimen (BGS GSM 85246; pl. 4, fig. 6) can be assigned to B.
sandbergeri without any doubts.
Bactroceras cf. sandbergeri (Barrande, 1867)
1994. Eobactrites sandbergeri (Barrande) - Kraft and Kraft, p. 9
(not figured).
1999. Bactroceras cf. sandbergeri - Marek, p. 413.
Material: Four specimens (MBHR 3228, 3229, 5359, 20279),
flattened fragments of phragmocones preserved as internal
moulds in green-grey shales.
Occurrence (Figs. 3, 4; Table 2): Prague Basin, central Bohemia,
Czech Republic; Ordovician, upper Dapingian, Klabava Formation,
Azygograptus-Tetragraptus Biozone: locality Klabava (Starý hrad).
Measurements: See Table 1.
Maximum shell Diameter of Diameter of
diameter (mm) shell/height siphuncle/Diameter
of chamber of shell
13.2 Living chamber Living chamber
4.5 1.5 1/11
45 2 to 3 1/7
25.3 2.9 1/8
Description: The studied material consists of parts of
orthoconic phragmocones, ranging in diameter from 5 to
12 mm. The angle of expansion ranges between 48 and 68. The
sutures are straight and directly transverse. Phragmocone
chambers are rather deep (height between 1/2 and 1/3 that of
the phragmocone diameter). The siphuncle is marginal and
tubular. Its diameter reaches 1/10 that of the phragmocone
diameter. Shell sculpture, where preserved, consists of straight and
dense, faint growth lines (7 growth lines per mm), directly
transverse to the longitudinal axis of the shell.
Remarks: Significant deformation of this material resulted in
the loss of diagnostic characters, making the assignment of the
specimens to B. sandbergeri uncertain. However, the siphuncle
position, apical angle, distance and course of sutures, as well as
course of growth lines are comparable to the structures described
in B. sandbergeri.
Bactroceras interpolatum (Barrande, 1867) nov. comb.
Fig. 9(A, B)
Partim. 1870. Bactrites Sandbergeri Barr. - Barrande, pl. 413, figs.
13-14.
1870. Orthoceras interpolatum Barr. - Barrande, pl. 417, figs. 3-5.
1999. ‘‘O.’’ interpolatum (Barrande) - Marek, pp. 413-416.
Holotype: specimen NM L 20994, figured by Barrande (1870:
pl. 413, figs. 13, 14), herein re-figured in Fig. 9(A).
Type locality and horizon: Králův Dvůr near Beroun,
central Bohemia; Upper Ordovician, upper Katian, Králův Dvůr
Formation.
Material: Two specimens, a living chamber NM L 13764
(Fig. 9(B)) from the locality Lejškov u Málkova (Fig. 3(11)) and a
part of phragmocone NM L 20994 (Fig. 9(A)) from the locality
Králův Dvůr (Fig. 3(12)), preserved in a carbonated nodule. The
specimen from Lejškov u Málkova locality is stratigraphically
confined to the lower part of the Králův Dvůr Formation, i.e., the
interval with Anticostia teres and Styracograptus lobatus. The
stratigraphic position of the specimen from Králův Dvůr locality
is not possible to determine precisely; it may correspond to the
interval with Anticostia teres and Styracograptus lobatus, or
Dicellograptus laticeps Taxon-range Zone. See Kraft et al. (2015)
for details on the biostratigraphy of the Králův Dvůr Formation, as
well as Fig. 6 and Table 2 herein.
Occurrence (Figs. 3 and 4): Prague Basin, Czech Republic;
Ordovician, upper Katian, Králův Dvůr Formation: localities
Lejškov u Málkova (interval with Anticostia teres and Styracograptus
lobatus) and Králův Dvůr (interval with Anticostia teres and
Styracograptus lobatus, or Dicellograptus laticeps Taxon-range
Zone).
Measurements: See Table 1.
Diagnosis: Bactroceras with straight, faint longitudinal ridges
on the surface, concavity of septa up to 30% and sub-marginal,
tubular siphuncle. Diameter of siphuncle about 1/10 of the
phragmocone diameter.
 M. Aubrechtová / Geobios 48 (2015) 193–211 205

Table 2 Table 2 (Continued )

Localities and stratigraphic occurrence of the studied specimens. Specimen Locality Graptolite biozonation
Specimen Locality Graptolite biozonation MBHR 5825 Osek u Rokycan C. retroflexus
MBHR 9535 Rokycany - Drahouš C. retroflexus
Bactroceras sandbergeri
MBHR 13239 Osek u Rokycan C. retroflexus
NM L 10331 Osek u Rokycan C. retroflexus
MBHR 15521 Osek u Rokycan C. retroflexus
NM L 20993 Osek u Rokycan C. retroflexus
MBHR 17825 Osek u Rokycan C. retroflexus
NM L 40988 Praha-Šárka C. retroflexus or D. clavulus
MBHR 20333 Osek u Rokycan C. retroflexus
NM L 40989 Mýto u Rokycan C. retroflexus
CGS PP545 Osek u Rokycan C. retroflexus
NM L 40990 Šárka-pole C. retroflexus or D. clavulus
CGS PP546 Osek u Rokycan C. retroflexus
NM L 40991 Šárka-pole C. retroflexus or D. clavulus
CGS PP547 Osek u Rokycan C. retroflexus
NM L 40992 Šárka-pole C. retroflexus or D. clavulus
CGS PP595 Osek u Rokycan C. retroflexus
NM L 40993 Šárka-pole C. retroflexus or D. clavulus
CGS CW11 Osek u Rokycan C. retroflexus
NM L 40994 Šárka-cihelna C. retroflexus or D. clavulus
CGS CW13 Osek u Rokycan C. retroflexus
NM L 40995 Šárka-pole C. retroflexus or D. clavulus
UK CHMHZ-OR-0001 Praha-Šárka C. retroflexus or D. clavulus
NM L 40996 Praha-Šárka C. retroflexus or D. clavulus
NM L 40997 Praha-Šárka C. retroflexus or D. clavulus Bactroceras cf. sandbergeri
NM L 40998 Praha-Šárka C. retroflexus or D. clavulus MBHR 3228 Klabava (Starý hrad) Azygograptus-Tetragraptus
NM L 40999 Praha-Šárka C. retroflexus or D. clavulus Biozone
NM L 41000 Praha-Šárka C. retroflexus or D. clavulus MBHR 3229 Klabava (Starý hrad) Azygograptus-Tetragraptus
NM L 41001 Kařı́zek C. retroflexus Biozone
NM L 41002 Cekov C. retroflexus MBHR 5359 Klabava (Starý hrad) Azygograptus-Tetragraptus
NM L 41003 Osek u Rokycan C. retroflexus Biozone
NM L 41004 Osek u Rokycan C. retroflexus MBHR 20279 Klabava (Starý hrad) Azygograptus-Tetragraptus
NM L 41005 Zbiroh C. retroflexus Biozone
NM L 41006 Mýto u Rokycan C. retroflexus
NM L 41007 Dı́ly u Rokycan C. retroflexus Bactroceras interpolatum nov. comb.
NM L 41008 Osek u Rokycan C. retroflexus NM L 13764 Lejškov u Málkova Interval with Anticostia teres
NM L 41009 Osek u Rokycan C. retroflexus and Styracograptus lobatus
NM L 41010 Mýto u Rokycan C. retroflexus (see Kraft et al., 2015)
NM L 41011 Šárka-cihelna C. retroflexus or D. clavulus NM L 20994b Králův Dvůr Interval with Anticostia teres
NM L 41012 Šárka-pole C. retroflexus or D. clavulus and Styracograptus lobatus
NM L 41013 Šárka-pole C. retroflexus or D. clavulus or Dicellograptus laticeps
NM L 41014 Šárka-cihelna C. retroflexus or D. clavulus Raxon-range Zone (see
NM L 41015 Šárka-cihelna C. retroflexus or D. clavulus Kraft et al., 2015)
NM L 41016 Šárka-cihelna C. retroflexus or D. clavulus
Bactroceras boliviensis nov. sp.
NM L 41017 Šárka-pole C. retroflexus or D. clavulus
UK UGP-O-0001b Chaupiuno, Bolivia B. minutus (P. densus)
NM L 41018 Zbiroh-Pětidomky C. retroflexus
UK UGP-O-0004 Chaupiuno, Bolivia B. minutus (P. densus)
NM L 41019 Praha-Šárka C. retroflexus or D. clavulus
a
NM L 41020 Praha-Šárka C. retroflexus or D. clavulus Lectotype.
b
NM L 41021 Zbiroh-Pětidomky C. retroflexus Holotype.
NM L 41022 Osek u Rokycan C. retroflexus
NM L 41023 Praha-Šárka C. retroflexus or D. clavulus
NM L 41024 Mýto u Rokycan C. retroflexus
NM L 41025 Osek u Rokycan C. retroflexus
NM L 41026 Osek u Rokycan C. retroflexus Description: The material consists of a portion of a
NM L 41027 Osek u Rokycan C. retroflexus phragmocone (Fig. 9(A)) and a body-chamber (Fig. 9(B)), with
NM L 41309 Zbiroh C. retroflexus
NM L 42945 unknown unknown
a maximum diameter of 13 mm and with a maximum height of
NM L 42946 Rokycany C. retroflexus 36 mm. The shell is straight, with a circular cross-section. The
NM L 42947 Praha-Šárka C. retroflexus or D. clavulus angle of expansion ranges between 48 and 68. The sutures are
NM L 42948 Praha-Šárka C. retroflexus or D. clavulus straight and transverse. 1.5 phragmocone chambers correspond
NM L 42949 Osek u Rokycan C. retroflexus
to the respective shell diameter; concavity is 1/3. The siphuncle
NM L 42950 Přı́lepy C. retroflexus or D. clavulus
NM L 42951 Úvaly C. retroflexus or D. clavulus is ventral, shifted from the shell wall. Cross-section of the
NM L 42952 Přı́lepy C. retroflexus or D. clavulus siphuncle is circular; its diameter equals 1/11 that of the
NM L 42953 Popovice C. retroflexus phragmocone diameter. The height of the preserved part of the
NM L 6576 Osek u Rokycan C. retroflexus living chamber (Fig. 9(B)) is 2–3 times the shell diameter. Shell
NM L 6577 Osek u Rokycan C. retroflexus
NM L 6578 Osek u Rokycan C. retroflexus
sculpture is not preserved, except faint, straight longitudinal
NM L 6579 Osek u Rokycan C. retroflexus ridges apparent on the surface of the living chamber (NM L
NM L 6580 Osek u Rokycan C. retroflexus 13764; Fig. 9(B)).
NM L 6581 Osek u Rokycan C. retroflexus Remarks: Shell morphology of specimen NM L 13764
NM L 6582 Osek u Rokycan C. retroflexus
(Fig. 9(B)), originally described as ‘‘Orthoceras’’ interpolatum
NM L 6583 Osek u Rokycan C. retroflexus
NM L 6584a Osek u Rokycan C. retroflexus Barrande, 1870, fully corresponds to the diagnosis of Bactroceras.
NM L 6585 Osek u Rokycan C. retroflexus The latter species is for this reason assigned to Bactroceras.
MBHR 1901 Rokycany C. retroflexus Specimen NM L 20994 (Fig. 9(A)), originally described as Bactrites
MBHR 1939 Rokycany C. retroflexus sandbergeri Barrande, 1870, is added to B. interpolatum nov. comb.,
MBHR 1951 Rokycany C. retroflexus
MBHR 1963 Osek u Rokycan C. retroflexus
based on its marked morphological similarities.
MBHR 1964 Osek u Rokycan C. retroflexus The high phragmocone chambers, sub-marginal siphuncle and
MBHR 1972 Osek u Rokycan C. retroflexus tubular connecting rings, as well as the concavity of the septa in
MBHR 1975 Osek u Rokycan C. retroflexus B. interpolatum nov. comb., fit well with the same features in B.
MBHR 1979 Osek u Rokycan C. retroflexus
angustisiphonatum. However, B. interpolatum nov. comb. exhibits
MBHR 1981 Osek u Rokycan C. retroflexus
MBHR 1983 Osek u Rokycan C. retroflexus longitudinal ridges on the surface of the shell, and the siphuncle
MBHR 1988 Osek u Rokycan C. retroflexus is narrower in diameter than in B. angustisiphonatum. Therefore,
MBHR 1993 Dı́ly u Rokycan C. retroflexus B. interpolatum nov. comb. is considered as a separate species
MBHR 5515 Rokycany C. retroflexus herein.
206 M. Aubrechtová / Geobios 48 (2015) 193–211
Fig. 10. Bivariate diagram showing variations in phragmocone proportional
chamber height during ontogeny in Bactroceras sandbergeri. Empty circles show
proportions of the embryonic shell of specimen NM L 10331; black circles show
proportions of mature specimens.
Bactroceras boliviensis nov. sp.
Fig. 6(A–D, G)
2000. Bactroceras aff. avus (Holm) - Marek et al., p. 56.
Derivation of the name: from ‘‘Bolivia’’, the country of the type
locality.
Holotype: The holotype specimen UK UGP-O-0001 is held at
the Faculty of Science, Charles University, Prague, and herein
figured in Figs. 6(A–D, G).
Type locality and horizon: Chaupiuno, near Tarija, Bolivia;
upper Lower Ordovician, upper Floian, Pircancha Formation,
Baltograptus minutus (Pseudophyllograptus densus) Biozone.
Material: Two specimens, UK UGP-O-0001 and UK UGP-O-
0004, were studied; these are slightly deformed phragmocones
preserved in grey-black micritic limestone. Both specimens were
Fig. 12. A. Latex cast of specimen NM L 40993 showing details of muscle scars. B. Idealized presumed outline of the muscle scars. Arrow points to aperture. Scale bars: 1 mm.
Fig. 11. Bivariate diagram showing variations in siphuncle proportional diameter
during ontogeny in Bactroceras sandbergeri. Empty circle shows proportions of the
embryonic shell of specimen NM L 10331; black circles show proportions of mature
specimens.
originally collected by Bernd Weber as part of project C3 of SFB
267, funded by the German Science Foundation.
Occurrence: Chaupiuno, near Tarija, Bolivia; upper Lower
Ordovician, upper Floian, Pircancha Formation: locality Chaupiuno
(Fig. 13).
Measurements: See Table 1.
Diagnosis: Bactroceras with a large shell diameter (45 mm),
transverse striae on the surface, and concavity of septa of 30%.
Siphuncle tubular, marginal. Diameter of siphuncle 1/8 to 1/7 of
the phragmocone diameter. Connecting rings very thin – 1/12 of
the diameter of the siphuncle.
Description: The holotype (UK UGP-O-0001; Fig. 6(A–D, G)) is
part of a phragmocone and living chamber, with a maximum
diameter of 45 mm. The shell is straight and circular in cross-
section. The angle of expansion is 58. The sutures are straight and
directly transverse. 2 to 3 phragmocone chambers correspond to
 M. Aubrechtová / Geobios 48 (2015) 193–211
Fig. 13. Palaeogeographic distribution of the genus Bactroceras during the Lower Ordovician. Map modified after Cocks and Torsvik (2006).
the respective shell diameter. The siphuncle (Fig. 6(G)) is ventral,
almost marginal, only very slightly removed from the shell wall.
Cross-section of the siphuncle is circular. Diameter of the siphuncle
is 1/7 of the phragmocone diameter. Septal necks are orthochoa-
nitic and closer (0.1 mm) to the shell wall than the connecting rings
(0.4 mm). Connecting rings are very thin, homogeneous and
tubular (or very slightly convex). Ultrastructure was not observed.
Thickness of the ring wall is 1/12 of the siphuncle diameter.
Sculpture consists of faint striae, inclined toward the longitudinal
axis of the shell. Primary cameral deposits were not observed.
Endosiphuncular deposits unknown.
Specimen UK UGP-O-0004 is part of a phragmocone, with a
maximum diameter of 25.3 mm. The shell is straight and circular in
cross-section. The sutures are straight, directly transverse. Three
phragmocone chambers correspond to the respective shell
diameter. The siphuncle is ventral, almost marginal, only very
slightly removed from the shell wall. Cross-section of the siphuncle
is circular. Diameter of the siphuncle is 1/8 of the phragmocone
diameter. Septal necks are orthochoanitic and closer to the shell
wall than the connecting rings. Connecting rings are very thin,
homogeneous and tubular. Ultrastructure was not observed.
Primary cameral deposits were not observed. Endosiphuncular
deposits are unknown.
4. Discussion
4.1. The genus Bactroceras and the bactritids
The systematic position of Bactroceras has long been a matter of
debate (see above, Section 1). If Bactroceras were a bactritid, then
the origin of this evolutionarily important group would be shifted
from the Early Devonian to the Early Ordovician. Similarities
between the two taxa include a slender, straight phragmocone with
a narrow, marginally situated siphuncle, and deep ventral lobe-like
structures on the ventral side of the shell (Erben in Moore, 1964).
Nevertheless, the present study supports the earlier conclusions of
Kröger and Mapes (2007) that the similarity of Bactroceras with
207
bactritids is only superficial, and more likely constitutes a
morphological convergence of two unrelated lineages caused by
a similar mode of life, rather than any evolutionary affinity. The key
argument used here relates to the external morphology of the
protoconch (Fig. 2(B)). The protoconch in Bactroceras is moderately
large, hemispherical, and possesses a small caecum and constric-
tion. In general morphology, it corresponds to protoconchs
previously described as Bactroceras angustisiphonatum (Evans,
2005: pl. 4, fig. 16) and other Early Ordovician orthocerid
cephalopods (Kröger, 2006). By contrast, remarkably constricted,
inflated embryonic chambers were described in bactritids, thus
displaying a significantly different morphology (Erben in Moore,
1964; Ruzhencev, 1962; Doguzhaeva, 1999; Kröger and Mapes,
2007). Additionally, Kröger and Mapes (2007) described a promi-
nent hyponomic sinus, lobes and saddles on both the sutures and
growth lines, as well as obliquely oriented septa in the bactritids.
Retractor muscle scars reported in the bactritids (Kröger and Mapes,
2004; Kröger et al., 2005) are dorsal, forming prominent,
moderately large lobes. Thus, at first glance the bactritids might
indeed resemble early orthocerids such as Bactroceras, but a more
detailed examination shows remarkable differences in the struc-
ture of their shells and organization of soft body. In agreement with
Evans (2005) and Kröger and Mapes (2007), ventral lobes on the
sutures, previously regarded as one of the main diagnostic features
of Eobactrites, are herein considered a consequence of taphonomic
processes. The close proximity of the connecting ring to the
phragmocone wall leaves no space for sediment, and consequently
a lobe-like structure appears on the internal mould (Evans, pers.
comm. 2014). Therefore the ventral lobes in Bactroceras are not
considered to have any systematic value.
4.2. Palaeobiogeographic and stratigraphic occurrence of the genus
Bactroceras
4.2.1. Lower Ordovician
No cephalopods are known from the Lower Ordovician
sediments of the Prague Basin (Marek, 1999). Benthic faunal
assemblages of Tremadocian age, however, point to the position of
208 M. Aubrechtová / Geobios 48 (2015) 193–211
Perunica near Baltica, within the temperate climatic zone
(Havlı́ček et al., 1994; Fatka and Mergl, 2009). Initiated during
the Floian, a transgression associated with the deepening of the
Prague basin facilitated the migration of new faunas to Perunica.
These faunas have affinities not only with Baltica, but also with
Eastern Avalonia and Armorica (Havlı́ček et al., 1994; Cocks, 2000;
Cocks and Torsvik, 2005, 2006; Fatka and Mergl, 2009) (Fig. 13).
During the Lower Ordovician, Bactroceras was restricted to
locations along the high-palaeolatitude margin of Gondwana
(Fig. 13), where it is found within morphologically monotonous
assemblages largely consisting of orthocerids or orthocerid-like
orthocones. Kröger and Evans (2011) described cephalopod
assemblages from the upper Tremadocian and lower Floian
sequences of the Montagne Noire, France. These cephalopod
assemblages consist of slender orthocones with thin to nearly
tubular, central or subcentral siphuncles. These cephalopods were
assigned to the families Eothinoceratidae and Rioceratidae
(Ellesmerocerida), Troedssonelidae (Dissidocerida), and Baltocer-
atidae (Orthocerida). Other groups include rare endocerids. So far,
Bactroceras and Annbactroceras mark the oldest known reports of
orthocerid cephalopods. Evans et al. (2013) reported the presence
of ?Bactroceras from the upper Tremadocian of Eastern Alborz, Iran.
The accompanying fauna additionally indicates a deeper shelf
environment, and a high latitude position of this region. General
morphology of cephalopods and the depositional environment of
the Montagne Noire show affinities with the Tremadocian/Floian
sequence of the Anti-Atlas, Morocco (Kröger and Lefebvre, 2012).
There, the cephalopod association, including Bactroceras, consists
exclusively of longiconic orthocones, although its taxonomic
composition is different. Also, cephalopods of upper Lower
Ordovician were investigated by Marek et al. (2000) in Chaupiuno,
Bolivia. That research showed the presence of Bathmoceras,
Protocycloceras, Bactroceras boliviensis nov. sp., and some undeter-
mined ellesmerocerids and endocerids. Last, Bactroceras was also
reported from the upper Tremadocian to middle Floian Seydisehir
Shales of the Taurus Mountains, southern Turkey (Dean and
Monod, 1970).
4.2.2. Middle Ordovician
The first cephalopods appear in the Prague Basin in the Middle
Ordovician (upper part of the Klabava Formation, i.e., uppermost
Dapingian; Fig. 4) (Kraft and Kraft, 1994). The sequence consists of
greenish to grey and reddish shales, tuffites and ferrolites, and
bears a low diversity cephalopod assemblage. Longiconic ortho-
cones prevail; however they are rare, and shells are mostly only
fragmentary and poorly preserved. The rare occurrence of
Bactroceras cf. sandbergeri (Orthocerida) is significant. The Baltic
species Rhynchorthoceras cf. angelini (Lituitida) and Bathmoceras
complexum (Ellesmerocerida) indicate relationships between
Baltic and Bohemian faunas during that time (Marek, 1999).
Benthic fauna shows affinities with Avalonia and Armorica
(Havlı́ček et al., 1994; Cocks, 2000; Cocks and Torsvik, 2005,
2006; Fatka and Mergl, 2009) (Fig. 5).
The Šárka Formation (lower to middle Darriwilian; Fig. 4)
represents a time interval during which a marked transgression
and deepening of the Prague basin took place, associated with
intensive basaltic volcanism. The fossil fauna is richly diverse and
mostly preserved within siliceous nodules; however, it is also
present in black shale lithofacies and ferrolites. Havlı́ček (1982)
and Havlı́ček and Vaněk (1990) assigned the assemblages to the
Euorthisina-Placoparia Community, which was interpreted as
inhabiting deep-water, soft-bottom environment situated beyond
the reach of wave and current activity. The presence of the
ichnogenus Arachnostega (Lefebvre, 2007) also indicates these
conditions. The Šárka Formation contains the most diversified
cephalopod fauna in the whole Ordovician of the Prague Basin
(Marek, 1999). The fossil record includes endocerids, actinocerids
(‘‘Orthoceras’’ bonum) and ellesmerocerids (Bathmoceras com-
plexum), but the most common elements are pseudorthocerids
and orthocerids (Mergl, 1978; Manda, 2008). The occurrence of
ellesmerocerids, actinocerids and endocerids suggests enhanced
faunal exchange between Perunica and Baltica (Marek, 1999). The
rather common presence of Bactroceras sandbergeri within the
Šárka Formation and the Darriwilian strata of Wales (Evans, 2005)
and Scandinavia (Holm, 1898; Sweet, 1958) might also suggest
faunal interchanges between Perunica, Avalonia and Baltica during
the Middle Ordovician (note, however, that many West-Gondwa-
nian cephalopod faunas are not yet sufficiently known and still
need to be described; Kröger, pers. comm. 2014). By contrast, the
benthic fauna (brachiopods, trilobites) of the Šárka Formation only
shows a weak similarity with Baltic assemblages, and rather
exhibits strengthening links with Armorica and other Western
Gondwana terranes (Havlı́ček et al., 1994; Fatka and Mergl, 2009).
The Šárka Formation and the succeeding Dobrotivá Formation
(upper Darriwilian to lower Sandbian; Fig. 4) are lithologically and
sedimentologically similar; however, the faunal composition
changes remarkably (Havlı́ček, 1998). According to Havlı́ček
et al. (1994) and Fatka and Mergl (2009), this may be related to
the Tornquist Sea functioning as a barrier against faunal migration
(see also Cocks, 2000; Cocks and Torsvik, 2005, 2006). According to
Havlı́ček et al. (1994) and Fatka and Mergl (2009), faunal migration
existed between Perunica and Armorica, as indicated by the
presence of a benthic fauna common to both regions. Cephalopod
communities are reduced in diversity, whilst lacking actinocerids
and endocerids. The only characteristic Baltic component is
Trocholites fugax (Babin and Gutiérrez-Marco, 1992), which marks
the first and only tarphycerid recorded from the Ordovician of the
Prague Basin (Manda, 2008). The majority of the cephalopod fauna
consists of orthocerids and pseudorthocerids which remain to be
revised. So far, Bactroceras has not been recorded.
During the Middle Ordovician, Bactroceras reached its maxi-
mum diversity and palaeobiogeographic distribution (Fig. 1). B.
sandbergeri occurs in Perunica, Baltica (Germany, Norway,
Sweden), and Avalonia (Wales). Another Middle Ordovician
species, B. angustisiphonatum, occurs in Baltica (Mecklenburg,
Germany; Rüdiger, 1889), South China (Zou, 1987), and Laurentia
(Nevada; Flower, 1968). Evans and King (1990) and Evans (2005)
also mention B. angustisiphonatum from the lower Dapingian of
Spitsbergen (Laurentia). B. huanghuaense, B. xianlingbuense, and B.
zhejiangense were reported from the Middle Ordovician of China by
Xu and Lai (1987) and Zou (1987).
4.2.3. Upper Ordovician
From the Upper Ordovician sequence of the Prague Basin, the
most diversified assemblage of cephalopods occurs within the
Králův Dvůr Formation (upper Katian to lowermost Hirnantian;
Fig. 4). The Formation consists of greenish claystones overlaid by a
layer of clayey carbonates and silty shales. The claystone sequence
is extremely poor in fossils, while a highly diversified fauna
appears within the carbonate sediments in the upper part of the
formation (Marek, 1952; Havlı́ček and Vaněk, 1966; Havlı́ček and
Mergl, 1982; Havlı́ček and Vaněk, 1990). The benthic fauna is
considered to reflect deep-water under low oxygen conditions
below the normal wave base. Similarities of faunal assemblages
between the Prague Basin, Baltica and Avalonia indicate that the
Tornquist Sea and Rheic Ocean were not serious barriers to the
migration of faunas at that time (Havlı́ček et al., 1994; Cocks,
2000; Cocks and Torsvik, 2005, 2006; Fatka and Mergl, 2009).
Cephalopods in the Králův Dvůr Formation are largely confined to
the argillaceous and carbonate sediments of the upper part of the
sequence (Marek, 1999). The taphocoenosis consists of members
of Pseudorthocerida and Orthocerida, including Bactroceras
 M. Aubrechtová / Geobios 48 (2015) 193–211
Fig. 14. Palaeogeographic distribution of the genus Bactroceras during the Upper Ordovician. Map modified after Cocks and Torsvik (2006).
interpolatum nov. comb., preserved within the clayey lithofacies.
Diestoceras primum also occurs, and marks the first appearance of
oncocerid cephalopods in the Prague Basin (Barrande, 1865-1874;
Marek, 1999) (Fig. 14).
Beyond the Prague Basin (Figs. 1, 14), Bactroceras subventrum
(Lai, 1987) and B. angustisiphonatum are recorded from the late
Sandbian sediments of South China. The latter species was also
reported from the lower Katian of New South Wales, Australia
(Glenister, 1952; Stait et al., 1985; Hewitt and Stait, 1985) and Irian
Jaya, Indonesia (Crick and Van Ufford, 1995).
4.3. Palaeobiogeographic and stratigraphic significance of the genus
Bactroceras
According to Kröger and Evans (2011), the palaeobiogeographic
and stratigraphic distribution of the earliest orthocerids implies an
origin of the order on the high-latitude margins of Gondwana
during the earliest Ordovician, their subsequent dispersion
towards lower, tropical latitudes starting with the Middle
Ordovician (see also Kröger, 2013). Kröger and Evans (2011)
stressed the typical characters of the Early Ordovician cephalopod
faunas of the high-palaeolatitude regions of Gondwana. During the
Early Ordovician (Tremadocian/lower Floian), assemblages con-
sisting of slender, longiconic orthocones were restricted to high-
palaeolatitude, temperate, cold-water regions, and were very
distinct from faunas of tropical regions where nautiloids and
endocerids dominated. According to Kröger and Evans (2011),
cephalopod assemblages of high-palaeolatitude regions started
expanding southwards in the lower Floian, continuing during the
Middle Ordovician, then becoming a common component of
presumably warm-water, low-palaeolatitude cephalopod commu-
nities. By contrast, elements of tropical faunas such as actinocerids
and tarphycerids only occasionally spread to high-palaeolatitude
locations (Marek, 1999; Manda, 2008). During the Upper Ordovi-
cian, orthocerids and orthocerid-like cephalopods were already a
common component of tropical faunas (Kröger, 2013).
During the Early Ordovician, Bactroceras was restricted to peri-
Gondwana (France, Iran, Turkey) and high-latitude regions of
209
Gondwana (Morocco, Bolivia) (Figs. 1, 13). During the Middle
Ordovician, the genus reached its maximum diversity; besides
Perunica and Avalonia, it was also found in Baltica, South China and
even as far as Laurentia (Figs. 1, 5). During the Upper Ordovician,
Bactroceras was restricted primarily to the low-latitude margin of
Gondwana (Australia, Indonesia) and South China with the
exception of Perunica, which marks its southernmost occurrence
(Figs. 1, 14). Such a palaeobiogeographic and stratigraphic
distribution of Bactroceras is in agreement with the views of
Kröger and Evans (2011), but additional material needs to be
studied. In particular the occurrences in China require to be revised
in order to obtain a more accurate picture of the expansion of
orthocerids during the Early Ordovician.
5. Conclusions
Bactroceras is a widespread Ordovician orthocerid cephalopod
(Figs. 1, 5, 13, 14). As the oldest known orthocerid, Bactroceras
(= Eobactrites) is of significant evolutionary importance; however,
only very limited, often poorly preserved material has been
previously available for study. With the exception of specimens
illustrated by Schindewolf (1932), a previously unstudied collec-
tion of more than one hundred rather well-preserved specimens
from the Prague Basin (Bohemia; Fig. 3) assigned to three species of
the genus Bactroceras has been examined in detail. The preserva-
tion of some of this material enables a better understanding of the
morphology of the shell, making possible a revision of the type
species of Eobactrites and Bactroceras in general. The stratigraphic
and palaeogeographic occurrences of Bactroceras have been
reevaluated. They might suggest an origin of the genus in high-
latitude, cold-water margins of Gondwana, and its subsequent
expansion to low-latitude, warm-water regions during the
Ordovician (see also Kröger and Evans, 2011).
Muscle scars, described for the first time in Bactroceras, show a
simple, unpaired and low dorsally-situated lobe (Figs. 2(G), 12).
Their morphology corresponds to ‘‘orthoceridan’’ muscle scars as
described by Mutvei (2002a,b), and especially muscle scars
illustrated by Mutvei (1957) for the genus Cochlioceras (family
210 M. Aubrechtová / Geobios 48 (2015) 193–211
Baltoceratidae). Based on the study of the morphology of juvenile,
as well as late ontogenetic and adult stages of B. sandbergeri, it can
be concluded that during ontogeny, the siphuncle diameter
declines (Fig. 11) and the phragmocone chambers shorten
(Fig. 10). The position of the siphuncle within the phragmocone
remains relatively unchanged during ontogeny, as does shell
sculpture, shape of the shell, and angle of expansion.
The species Bactroceras avus Holm, 1898 is herein placed in
synonymy with Bactroceras sandbergeri (Barrande, 1867), based on
their strong morphological similarities (mainly proportions of the
shell and position of the siphuncle) as well as the corresponding
stratigraphic occurrence of both species. A new species, Bactroceras
boliviensis from the upper Floian of Chaupiuno, Bolivia, is erected. It
is based on two specimens previously mentioned by Marek et al.
(2000). The species differs from other species of the genus mainly
in having a remarkably large phragmocone diameter. Orthoceras
interpolatum Barrande, 1870 from the Králův Dvůr Formation,
Bohemia (Katian), is here assigned to Bactroceras. B. interpolatum
nov. comb. and B. angustisiphonatum from the Upper Ordovician of
Australia and Indonesia are the stratigraphically youngest species
of Bactroceras. The assignment of Orthoceras gossei Etheridge, 1893
to Bactroceras (Teichert and Glenister, 1952) is rejected. The type
specimen of O. gossei illustrated by Etheridge (1893: pl. 1, figs. 9,
10) differs from Bactroceras in having extremely short phragmo-
cone chambers. While Cochlioceras and closely related genera
display a relatively wide siphuncle with endosiphuncular deposits
and rather short phragmocone chambers, Bactroceras, Annbactro-
ceras and Eosomichelinoceras are characterized by relatively higher
phragmocone chambers and a very narrow siphuncle without
endosiphuncular deposits. For these reasons, the present author
believes that future detailed studies of the morphology of these
cephalopods are needed to show whether the Baltoceratidae
constitute a monophyletic taxon or is in fact paraphyletic or
polyphyletic.
Acknowledgements
I especially thank Š. Manda (Czech Geological Survey, Prague)
for helpful advices concerning the morphology, evolution and
ecology of orthocerids. Deeply acknowledged is V. Turek (National
Museum, Prague), who made accessible the collection of Bactro-
ceras at the National Museum, provided me with photographs of
the Swedish specimen of B. avus, and helped with the corrections of
the manuscript. I thank B. Kröger (Finnish Museum of Natural
History, Helsinki, Finland) and D.H. Evans (Natural England,
Peterborough, England, United Kingdom) for kindly providing
some of the important papers and information on the species of the
studied genus. I acknowledge the reviewers, B. Kröger, D. H. Evans
and K. Histon (University of Modena and Reggio Emilia, Modena,
Italy), who helped to greatly improve the manuscript with their
comments and suggestions. Very special thanks belong to F.
Schönian (Humboldt Universität, Berlin), B. Weber (Freie Uni-
versität, Berlin) and J. Marek (Faculty of Science, Charles
University, Prague) who provided me with specimens of Bactro-
ceras from Bolivia, as well as important information related to
geology and stratigraphy of the Ordovician of southern Bolivia.
Many thanks are due to P. Budil (Czech Geological Survey, Prague)
and M. Polechová (Czech Geological Survey, Prague) for enabling
the study of specimens housed at the Czech Geological Survey in
Prague. I acknowledge Petr Kraft (Faculty of Science, Charles
University, Prague) for advice on the biostratigraphy of the
Ordovician of the Prague Basin. I thank M. Mergl (University of
West Bohemia) for the help in studying and photographing of
specimens housed at the Dr. Bohuslav Horák Museum in Rokycany.
Special thanks are due to P. Daneš for making English language
corrections, and M. Valent for essential help with photographs and
figures. Finally, I thank V. Kozák for kindly providing well-
preserved specimens of the studied genus.
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