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, JUNE 2011, 62, 3, 203—209 doi: 10.2478/v10096-011-0017-8
First record of the genus Bronnothyris (Brachiopoda:
Megathyrididae) from the Oligocene of the Mainz Basin
(Germany)
MARIA ALEKSANDRA BITNER
1
and ANDREAS KROH
2
1
Institute of Paleobiology, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warszawa, Poland; bitner@twarda.pan.pl
2
Natural History Museum Vienna, Department of Geology & Paleontology, Burgring 7, 1010 Vienna, Austria; andreas.kroh@nhm-wien.ac.at
(Manuscript received July 9, 2010; accepted in revised form October 13, 2010)
Abstract: The genus Bronnothyris, with the type species Terebratula bronnii Roemer, 1841, was erected for those
Argyrotheca species that have septal flanges extended ventrally from the dorsal valve. Four other Late Cretaceous and
one Early Paleocene species were attributed to this genus (i.e. Argyrotheca coniuncta Steinich, 1965, A. lacunosa
Steinich, 1965, A. obstinata Steinich, 1965, A. stevensis Nielsen, 1928 and A. rugicosta Zelinskaya, 1975). After exami-
nation of Oligocene material from Waldböckelheim, Mainz Basin, Germany we transfer the species Argiope subradiata
Sandberger, 1862 into the genus Bronnothyris. This new combination extends the stratigraphic range of Bronnothyris
into the Oligocene.
Key words: Oligocene, Germany, Mainz Basin, Brachiopoda, Megathyrididae, Bronnothyris.
Introduction
The genus Bronnothyris, with the type species Terebratula
bronnii Roemer, 1841, was erected by Popiel-Barczyk &
Smirnova (1978) for Late Cretaceous megathyridid brachio-
pods resembling Argyrotheca but having short septal flanges
that extend ventrally from the dorsal septum. Apart from
Argyrotheca bronnii, Popiel-Barczyk & Smirnova (1978) at-
tributed to this genus four other species from the Upper Creta-
ceous white chalk: A. coniuncta Steinich, 1965, A. lacunosa
Steinich, 1965, A. obstinata Steinich, 1965 and A. stevensis
Nielsen, 1928. Later one further species, Argyrotheca rugicosta
Zelinskaya, 1975 from the Lower Paleocene of Ukraine was
transferred to Bronnothyris by Smirnova et al. (1983).
In the present study we have placed another Argyrotheca
species, A. subradiata (Sandberger, 1862) from the Oligo-
cene of Germany in the genus Bronnothyris, thus consider-
ably extending the stratigraphic range of the genus. The aim
of this paper is to re-describe and re-illustrate this poorly
known species.
Geological setting and material
The Mainz Basin lies along the Rhine Graben fault sys-
tem in south-western Germany. It is subdivided into a
northern part (Rüsselsheim Basin) and a southern part
(Eisenberg Subbasin, Marnheim Bay). The material studied
here comes from the Rüsselsheim Basin. There the sedi-
mentary succession started in the Middle Eocene with the
deposition of terrestrial and limnic basal clays. These un-
fossiliferous beds locally extend into the Late Eocene and
are not exposed on the surface, being known only from
wells. In the Priabonian to Lower Rupelian limnic and flu-
vio-marine beds of the Pechelbonn Group were deposited.
Earlier interpreted as basal beds and/or marginal facies of
the “Rupelton” (Bodenheim Formation), these deposits are
now regarded as separate lithostratigraphic units. The
Pechelbonn Group contains a rich fauna and flora deriving
from a largely limnic to brackish shallow-water setting.
Above the Pechelbonn Group, the Rupelian to Chattian
Selztal Group is developed and it is these units that have
delivered the marine fossils for which the Mainz Basin is
famous. The Selztal Group is subdivided into several for-
mations, starting with the laterally interfingering Boden-
heim (former “Rupelton”) and Alzey Formation (former
“Meeressand”), overlain by the Stadecken Formation
(former “Schleichsand” and “Schleichsandmergel”), which
in turn is overlain by the Sulzheim Formation (former
“Cyrenenmergel”, “Cyrenenkalk”, and “Süßwasserschich-
ten”). On top of these beds, the Mainz Group with a rich
succession of carbonate platform deposits interfingering
with siliciclastic deposits of the Rhine Graben is devel-
oped. These beds are of Late Oligocene (Chattian) to Early
Miocene (Burdigalian) age. For more detailed data on the
lithostratigraphic subdivision of the Mainz Basin and asso-
ciated biostratigraphic data see Grimm (2002, 2005) and
Grimm & Grimm (2003), which also contain an extensive
description of the outcrops in the area, both historical and
modern.
The material studied here derives from the Alzey Formation
exposed in the area of Waldböckelheim, 10 km west of Bad
Kreuznach, SW Mainz, Germany (Fig. 1). There, the Alzey
Formation crops out on the southern and south-western slopes
of the Welschberg, north of Waldböckelheim and on the
north-western slope of the Heimberg, south-east of
Waldböckelheim. Both areas are classical localities of the 19
th
century and have delivered a huge number of fossils (predomi-
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nantly mollusks), both in number of species and
specimens. Grimm & Grimm (2003) attribute the
exceptional high diversity of the Waldböckelheim
deposits to the fact that these faunas actually derive
from a variety of small sandpits that exposed dif-
ferent levels in the succession and possibly stem
from slightly different paleoenvironments. Like
most Waldböckelheim material, the specimens
studied here carry no specific sandpit or sub-locali-
ty information. It seems likely, though, that they
derive from the outcrops at Heimberg which are
characterized by their abundant occurrence of
Argyrotheca subradiata (Grimm & Grimm 2003:
p. 91). According to Grimm et al. (2000) the Alzey
Formation was deposited during the Rupelian,
based on calcareous nannoplankton data (Zones
NP23 and lower NP24).
The material is housed in the Institute of Paleobio-
logy, Warszawa under the number ZPAL Bp.67 and
in the Natural History Museum Vienna under the
numbers NHMW 1863/0017/0061, 1866/0058/0068,
1868/0001/0820, and 2010/0169/0001 to …/0007.
cene): 4 complete specimens, 2 ventral valves, 12 dorsal
valves (ZPAL Bp.67/1—4, NHMW 2010/0169/0001 to
.../0007); additionally numerous complete and disarticulated
specimens are kept at the NHMW under the nos. NHMW
1863/0017/0061, 1866/0058/0068, and 1868/0001/0820.
D i m e n s i o n s (in mm):
Specimen number Length Width Thickness
ZPAL Bp.67/2
5.2 5.8 3.4
ZPAL Bp.67/3 (neotype)
6.0 5.7 3.4
D e s c r i p t i o n: Shell small, subrectangular to subpentago-
nal in outline, usually wider than long, ventriconvex. Shell
surface covered with up to 10 rounded ribs that become less
distinct at the anterior margin; growth lines distinct. Both
valves coarsely punctate. Beak high, suberect to erect, eroded
to some degree in most specimens (Fig. 2E,H), suggesting a
very short pedicle and close attachment to the substrate.
Sometimes part of the dorsal valve is also corroded (Fig. 2H).
Beak ridges sharp, subtending an angle of about 90—100° at
umbo. Interarea relatively narrow, transversely striated. Fora-
men large, subtriangular, hypothyrid, flanked by two narrow,
disjunct deltidial plates. Hinge line long, straight, often equal
to maximum width. Lateral commissures straight, anterior
commissure rectimarginate.
Ventral valve interior with short wide teeth lying parallel to
hinge line; teeth bear weak diagonal ridges. Pedicle collar
wide, supported by a slender median septum that extends to
about mid-valve; anteriorly to septum there are two or three
shallow ovoid depressions to accommodate serrations of the
dorsal septum.
Dorsal valve nearly flat, its interior bears short, widely di-
vergent inner socket ridges. Cardinal process in form of
short, indistinct ridges, situated between socket ridges.
Hinge plates broad, fused mid-dorsally to form a single co-
Fig. 1. Geographic position of Waldböckelheim in the Mainz Basin. Inset
shows position of map in relation to Germany. Distribution of Lower Oligo-
cene marine deposits (Alzey Formation, Bodenheim Formation) based on
Grimm & Steurbaut (2001: fig. 1C).
Systematics
Phylum: Brachiopoda Duméril, 1806
Subphylum: Rhynchonelliformea Williams, Carlson,
Brunton, Holmer & Popov, 1996
Class: Rhynchonellata Williams, Carlson, Brunton, Holmer
& Popov, 1996
Order: Terebratulida Waagen, 1883
Superfamily: Megathyridoidea Dall, 1870
Family: Megathyrididae Dall, 1870
Genus: Bronnothyris Popiel-Barczyk & Smirnova, 1978
T y p e s p e c i e s: Terebratula bronnii Roemer, 1841 by orig-
inal designation of Popiel-Barczyk & Smirnova (1978: 41).
Bronnothyris subradiata (Sandberger, 1862)
Figs. 2—4
?p.p. 1853 Terebratula sp. – Sandberger, p. 8
*1862 Argiope subradiata Sandberger, pl. 34, fig. 4, 4a—d
1863 Argiope subradiata Sand. – Sandberger, p. 386—387
1883 Argiope subradiata Sandberger – Lepsius, p. 56
1986 Argyrotheca subradiata (Sandberger) – von der Hocht, p. 208
T y p e m a t e r i a l: The syntypes studied by Sandberger
(1863) came from the Weinkauff collection from a locality at
Waldböckelheim-Heimberg. According to K. Grimm from the
Natural History Museum Mainz (pers. comm. Dec. 2007), the
Weinkauff collection was kept at the Bayrische Staatssamlung
für Geologie und Paläontologie, Munich, where it was de-
stroyed during the Second World War (Gürs 1995). Thus, we
have selected a neotype for this species.
N e o t y p e: The specimen figured in Fig. 2D—G, ZPAL
Bp.67/3.
M a t e r i a l e x a m i n e d: Waldböckelheim, Rheinland-
Pfalz, Germany (Alzey Formation, Rupelian, Early Oligo-
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herent platform; the boundary between inner and outer plates
are indistinct, indicated only by a shallow fold. Crura very
short; crural processes medianly directed, massive and rela-
tively long (Figs. 2I, 3H). Descending branches curved later-
ally and united with valve floor; they emerge from the valve
floor and are attached to the median septum. Short septal
flanges extend ventrally from dorsal septum (Figs. 2G,
4D—F). Septum high, triangular in profile, beginning from
hinge plates and sloping towards the anterior margin with
3—4 serrations.
The secondary fibres of the shell observed on the inner
surface are regularly packed forming a characteristic mosaic
(Fig. 4C).
R e m a r k s: The presence of the septal flanges which ex-
tend ventrally from the dorsal septum clearly indicates the
attribution of the investigated specimens to the genus
Bronnothyris. Externally, in shell outline and number of ribs,
the specimens are most similar to the Late Cretaceous B.
bronnii (Roemer, 1841), from which they differ internally,
however; B. bronnii possesses hinge plates that are devel-
oped as two concave circular separated discs, situated anteri-
orly to the cardinal process (Steinich 1965; Popiel-Barczyk
1968; Surlyk 1972, 1982; Bitner & Pisera 1979; Johansen
1987; Johansen & Surlyk 1990).
In turn, B. coniuncta (Steinich, 1965) differs externally
from B. subradiata in having strongly transversely elongate
Fig. 2. A—I – Bronnothyris subradiata (Sandberger, 1862), Oligocene, Waldböckelheim, Mainz Basin, Germany. A – dorsal view of
complete specimen, ZPAL Bp.67/1; B, C – outer and inner views of ventral valve, ZPAL Bp.67/2; D—G – ventral, dorsal, lateral and pos-
terior views of complete specimen, septal flanges extending from a dorsal septum arrowed, neotype, ZPAL Bp.67/3; H, I – dorsal and pos-
terior views of complete specimen, massive crural processes visible (I), ZPAL Bp67/4. All SEM.
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Fig. 3. A—I – Bronnothyris subradiata (Sandberger, 1862), Oligocene, Waldböckelheim, Mainz Basin, Germany. A – Ventral valve, ex-
terior view, NHMW 2010/0169/0007; B – ventral valve, inner view, NHMW 2010/0169/0005; C – dorsal valve, inner view, NHMW
2010/0169/0006; D – dorsal valve, lateral view, NHMW 2010/0169/0002; E, F – dorsal valve, inner and lateral views, NHMW 2010/
0169/0001; G – inner view of dorsal valve, NHMW 2010/0169/0004; H, I – dorsal valve, inner and lateral views, NHMW 2010/0169/
0003. All SEM.
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Fig. 4. A—F – Bronnothyris subradiata (Sandberger, 1862), Oligocene, Waldböckelheim, Mainz Basin, Germany. A, B – lateral views of
dorsal valve interior to show median septum and cardinalia, NHMW 2010/0169/0003, NHMW 2010/0169/0006; C – inner surface of ven-
tral valve, visible punctae and mosaic of secondary fibres, NHMW 2010/0169/0005; D – dorsal valve, side view of median septum,
NHMW 2010/0169/0001; E, F – interior views of dorsal valves to show details of septal flanges extending from median septum, NHMW
2010/0169/0003, NHMW 2010/0169/0001. All SEM.
outline, although its hinge plates, as in B. subradiata, are
fused and form a coherent, distinct platform (Steinich 1965;
Bitner & Pisera 1979; Surlyk 1982; Johansen 1987; Johansen
& Surlyk 1990).
The species B. obstinata (Steinich, 1965) and B. lacunosa
(Steinich, 1965) are distinguished from the Oligocene B. subra-
diata by having a smaller shell size and fewer ribs (Steinich
1965). Additionally, hinge plates in B. lacunosa form two
concave discs, whereas in B. obstinata hinge plates form a sin-
gle plate which is much narrower than that in B. subradiata.
The Oligocene specimens can also be easily distinguished
from the fifth Late Cretaceous species, B. stevensis (Nielsen,
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1928) which has disc-shaped, concave hinge plates (Johansen
1987; Simon 1998). Additionally, two septal flanges are
higher in B. stevensis than in B. subradiata (see Simon 1998:
pl. 7, fig. 4).
From the Early Paleocene B. rugicosta (Zelinskaya, 1975),
the investigated specimens differ strongly in ornamentation;
in B. rugicosta the shell surface is coarsely ribbed (Zelinskaya
1975: pl. 16, figs. 3, 4a, 5, 6a).
Argyrotheca wansinensis Vincent, 1923 from Wansin,
southern Belgium (now attributed to the early—middle Thane-
tian; compare De Geyter et al. 2006: p. 205; Jagt et al. 2007:
p. 40) is quite similar to A. subradiata. Vincent’s (1923) illus-
trations, however, are insufficient for proper comparison. We
were unable to trace his type material, nor did we succeed in
locating topotypic specimens. Thus for the time being, the re-
lation between these two forms, albeit similar, must remain
unresolved.
Bitner & Schneider (2009) have described Late Burdigalian
(Early Miocene) Argyrotheca cf. subradiata (Sandberger,
1862) from the Bavarian part of the Molasse Basin (Para-
tethys). Externally that material is similar to that from the
Mainz Basin, but differs in having much narrower hinge
plates, which are very wide in typical B. subradiata and form
a single plate. A. cf. subradiata also lacks the septal flanges
extending from the septum (Bitner & Schneider 2009) which
prevents the attribution of this species to Bronnothyris.
Externally the studied specimens also resemble Megathiris
detruncata (Gmelin, 1791), however, they can be readily dis-
tinguished internally by the lack of the lateral septa (e.g.
Logan 1979; Bitner 1990).
O c c u r r e n c e: Early Oligocene (Rupelian) of Wald-
böckelheim, Mainz Basin, Germany. This species has also
been reported (Sandberger 1863; Lepsius 1883; von der
Hocht 1986) from numerous other sites in the Mainz Basin
(e.g. Eckelsheim, Alzey-Weinheim, and Heimberg). Addi-
tionally, numerous “Argyrotheca” specimens from these and
further localities are available at the Natural History Museum
of Mainz (Grimm & Grimm 2005), which need to be studied
in future to confirm their specific identity with Bronnothyris
subradiata.
Discussion
Untill recently, the family Megathyrididae contained two
extant (Megathiris, Argyrotheca) and two fossil (Bronnothy-
ris, Phragmothyris) genera. Recently Álvarez et al. (2008b)
erected a new genus Joania (with type species Terebratula
cordata Risso, 1826) for those Argyrotheca species that have
a narrow hinge line, prominent cardinal process and tubercles
on the inner margin. Apart from Phragmothyris, all the genera
are externally very similar; the genus Phragmothyris from the
Eocene of Cuba, in contrast, differs strongly from other mem-
bers of the family in having a multicostellate surface (up to 40
ribs) and conjunct deltidial plates forming a symphytium
(Cooper 1955).
The placement of the Oligocene species Argiope subradiata
in the genus Bronnothyris considerably extends the strati-
graphic range of this genus. So far Bronnothyris has been re-
corded from the Upper Cretaceous to Lower Paleocene (Lee et
al. 2006). Thus, the genus Bronnothyris presently contains 7
valid species: B. bronnii, B. coniuncta, B. lacunosa, B. obsti-
nata, B. stevensis, B. rugicosta and B. subradiata. Judging
from the data available, the genus seems to have a distribution
restricted to Europe.
In adult specimens of A. cuneata (Risso, 1826) two ventro-
laterally directed prongs often diverge from the septum poste-
rior to the crest (Logan 1979; Brunton 1988; Álvarez et al.
2008a,b); they are interpreted as possible rudimentary ascend-
ing branches. It is possible that the septal flanges extending
from the septum in Bronnothyris could also be interpreted as
rudimentary elements of ascending branches.
The extant species Argyrotheca schrammi (Crosse & Fischer,
1866), from Barbados illustrated by Álvarez et al. (2008b:
fig. 7C,D) exhibits two short septal flanges as well. Although
further data are needed to check the consistency of this feature
in A. schrammi, this suggests that this species might also belong
to Bronnothyris, further extending the range of the genus, both
in terms of stratigraphic and spatial distribution. A. schrammi
differs from B. subradiata by the presence of spines on the pos-
terior slope of the median septum.
Argyrotheca is a very diverse genus, both today and in the
Cenozoic, and even back to the Cretaceous (Cooper 1977).
Further detailed examinations of other Cenozoic and extant
Argyrotheca species may reveal that there are more, as yet
unrecognized species of Bronnothyris. In the Treatise Lee et
al. (2006) suggest, that “Argyrotheca may include species
that should be assigned to several different genera”. Thus,
further investigations and a thorough revision are needed,
and the diversity of Bronnothyris may increase significantly
with future studies.
Acknowledgments: This research was supported by the EU
Grant, SYNTHESYS Project AT-TAF-2610 and the Grant
No. N N307 129837 from the Ministry of Science and Higher
Education (Poland) to MAB. We thank Kirsten and Matthias
Grimm for providing literature on the Mainz Basin and infor-
mation regarding the whereabouts of Sandberger’s type mate-
rial. We gratefully acknowledge the two referees Daphne E.
Lee (Univerity of Otago, Dunedin) and Kirsten Grimm (Natu-
ral History Museum Mainz) for their valuable reviews. The
SEM photos were taken in the SEM laboratory of the Institute
of Paleobiology (Warszawa) using a scanning microscope
Philips XL-20 and at the Natural History Museum Vienna, us-
ing a Jeol 6400 scanning microscope.
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