Cystoporate bryozoans from the Lower - Middle Devonian of NW Spain

Abstract

Ten bryozoan species belonging to the Order Cystoporata are described from the Lower Devonian of the Cantabrian Mountains, NW Spain. Three genera, with one species respectively, are new: Physallidopora cantabrica gen. nov. sp. nov., Dolichozoon ramosum gen. nov. sp. nov. and Hiberotrypa dubia gen. nov. sp. nov. Two additional species are new: Fistuliporidra triangulata sp. nov. and Fistuliporidra hibera sp. nov. The studied fauna shows palaeobiogeographic relations to the Czech Republic, Germany, Poland and North America.

Full text

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© 2007 E. Schweizerbart’sche Verlagsbuchhandlung, D–70176 Stuttgart
Paläontologische Zeitschrift
2007, Vol.
81/2
, p. 113–117, 30-06-2007
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A cystoporate bryozoan species from the Zechstein (Late Permian)
A
NDREJ
E
RNST
, Kiel
with 1 figure and 1 table
E
RNST
, A. 2007. A cystoporate bryozoan species from the Zechstein (Late Permian). – Paläontologische Zeitschrift
81
(2): 113–117, 1 fig., 1 tab., Stuttgart, 30. 6. 2007.
Abstract:
The bryozoan species
Choniopora radiata
S
CHAUROTH
, 1854 is recognised as
Fistulipora radiata
(S
CHAU-
ROTH
, 1854). New material from the Lower Zechsteinkalk (Late Permian) of Rügen island and Thuringia allows a
comprehensive description of its interior morphology. This is the first unequivocal record of a cystoporate bryozoan
from the Zechstein.
Keywords:
Bryozoa • Zechstein • Late Permian • taxonomy • morphology • palaeobiogeography
Kurzfassung:
Die Bryozoenart
Choniopora radiata
S
CHAUROTH
, 1854 wurde als
Fistulipora radiata
(S
CHAUROTH
,
1854) bestimmt. Neue Proben aus dem Unteren Zechsteinkalk der Insel Rügen und aus Thüringen erlauben eine um-
fassende Beschreibung der inneren Morphologie dieser Art. Dies ist der erste Nachweis einer cystoporaten Bryozoe
aus dem Zechstein.
Schlüsselwörter:
Bryozoa • Zechstein • Ober-Perm • Taxonomie • Morphologie • Paläobiogeographie
Introduction
The fauna of the Zechstein (Late Permian) has been
studied for more than 200 years. However, a large
number of existing publications reveal many uncertain
descriptions which have produced numerous taxonomic
problems. A closer investigation of known and new lo-
calities and collections of Zechstein fossils allows to
solve existing uncertainties and to make interesting con-
clusions.
Comprehensive studies of bryozoans from the Zech-
stein (e.g. K
ORN
1930; D
REYER
1961; S
OUTHWOOD
1985, 1990; most recently E
RNST
2001) have revealed
(Ca 1) only a reduced diversity of the bryozoan fauna
from reefs and bedded facies of the Zechsteinkalk.
Among all the bryozoan taxa, fenestrates are dominating
in assemblages of the Zechstein. Trepostomes are less
abundant and diverse, but a common group, characteristic
mostly for the bedded facies. A single cyclostome bryo-
zoan,
Corynotrypa voigtiana
(K
ING
, 1850) is also known
from the Zechstein. However, cystoporate bryozoans, al-
though abundant elsewhere during the Permian, were not
yet reported from the Zechstein. A cystoporate species
Coscinotrypa dubia
(G
EINITZ
, 1848), mentioned by M
O-
ROZOVA
(1970: 276) and later introduced in many other
publications, does not belong to this genus, but to an in-
determinable trepostome bryozoan (E
RNST
2001: 137).
In his publication from the year 1854, Carl Friedrich
von Schauroth described the new genus and species
Cho-
niopora radiata
from the Lower Zechstein of Thieschitz
near Gera in Thuringia. He assigned it as new bryozoan
species and genus having similarities with celleporids
(Cheilostomata). This fossil, however, was not recog-
nized as a bryozoan in later students. It was cited as a pos-
sible synonym of the brachiopod genus
Acanthocrania
W
ILLIAMS
, 1943 (A
GER
et al. in M
OORE
1965: H290).
However, a restudy of collections from the Bundesanstalt
für Geowissenschaften und Rohstoffe (BGR), Berlin, and
from the Museum für Naturkunde in Gera, revealed some
specimens of a cystoporate bryozoan, which belong to the
genus
Fistulipora
M’C
OY
, 1849. This bryozoan is super-
ficially very similar to
Choniopora radiata
S
CHAUROTH
,
1854. Certain details from the original description such as
a radial arrangement of zooecia, presence of horse-shoe
shaped lunaria (“ovarial pore at each aperture”) and a ve-
sicular skeleton between autozooecia (“mass with small
pores”) support the assumption that this fossil belongs to
cystoporate bryozoans. The description of S
CHAUROTH
Address of the author: Andrej Ernst, Institut für Geowissenschaften, Christian-Albrechts-Universität, Ludewig-Meyn-Straße 10,
D-24118 Kiel; e-mail <ae@gpi.uni-kiel.de>.

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(1854) is based on a single specimen. Unfortunately, this
specimen could not be traced in the collection of C.F.
VON
S
CHAUROTH
, housed at the Naturkunde-Museum Coburg
(E
CKHARD
M
ÖNNIG
, pers. comm.). From the newly dis-
covered material, a neotype and hypotypes are chosen.
Material and methods
The present study is based on two collections of Zech-
stein rocks and fossils. The collection of the Bundes-
anstalt für Geowissenschaften und Rohstoffe (BGR),
Berlin, contains drilling cores from the borehole Samtens
101, taken on the island of Rügen. Zechstein rocks were
investigated from cores 1736.86–1739.35 m, and
1744.05–1747.7 m. From this material 35 standard thin
sections were produced. Bryozoans are embedded in
grainstones, partly wackestones, which contain abundant
bryozoans, foraminifers, brachiopods, gastropods, bi-
valves, as well as crinoid fragments and echinoid spines.
The rocks are partly dolomitized and contain gypsum in-
tercalations and inclusions. The bryozoan assemblage
represents a typical association of the bedded facies of the
Lower Zechsteinkalk, depositional cycle Ca 1. Drilling
core material and thin sections are deposited at the Bun-
desanstalt für Geowissenschaften und Rohstoffe (BGR),
Berlin, the neotype and additional figured specimen un-
der collection numbers X 12896 and X 12897.
Further material comes from the collection of the
Museum für Naturkunde in Gera. Four brachiopods
(
Horridonia horrida
) from the collection of B
RET-
SCHNEIDER
(1959) (B
RETSCHNEIDER
is the maiden
name of Eva D
REYER
, who published her results in
1961) contained discoid colonies of the new species.
They were partly misidentified as holding organs of
Protoretepora ehrenbergii
(G
EINITZ
, 1861), partly as
“bryozoan holding organs of unknown origin”. Indeed,
specimens contain also some holding discs of
Kingopo-
ra ehrenbergii
(G
EINITZ
, 1861). Furthermore, brachio-
pod shells are incrusted by the trepostome bryozoan
Dyscritella microstoma
E
RNST
, 2001. A single acetate
peel has been produced by the author from the polished
surface of a sample III/C2319, one of the two paratypes.
According to the available labels this material was sam-
pled from the Lower Zechsteinkalk at the localities
Röpsen (III/C2627) and Roschütz (III/C2319 and III/
C2615) near Gera. The investigated material is housed
at the Museum für Naturkunde in Gera.
Systematic palaeontology
Phylum Bryozoa E
HRENBERG
, 1831
Order Cystoporata A
STROVA
, 1964
Family Fistuliporidae U
LRICH
, 1882
Genus
Fistulipora
M’C
OY
, 1850
Type species
:
Fistulipora minor
M’C
OY
1850 [=
F. incrus-
tans
(P
HILLIPS
, 1836)]; Lower Carboniferous, England.
Diagnosis
: Massive, encrusting or ramose colonies. Cy-
lindrical autozooecia with thin walls and complete dia-
phragms. Apertures with a lunarium. Autozooecia sepa-
rated by the extrazooidal vesicular skeleton. Maculae
usually developed.
Occurrence
: Ordovician – Permian; worldwide.
Fistulipora radiata
(S
CHAUROTH
, 1854)
Fig. 1; Tab. 1
* 1854
Choniopora radiata
S
CHAUROTH
: 564, pl. 20 fig. 7.
? 1861
Stenopora columnaris
var.
incrustans
. – G
EINITZ
:
pl. 21 figs. 5, 6.
Holotype
: (? missing) figured by S
CHAUROTH
1854, pl. 20
figs. 7a–c.
Neotype
:
III/C2319 (two colonies and one acetate peel).
Hypotypes
: X 12896 and X 12897, three thin sections (bore-
hole Samtens 101; 1736.86–1739.35 m).
Nomenclatural remarks
: The neotype was chosen according
articles 75.3.4 and 75.3.6 of the International Code of Zoolo-
gical Nomenclature (ICZN). The holotype based on a single
specimen is missing (see Introduction). Type locality (Thie-
schitz near Gera, Thuringia) is not accessible. The neotype
was chosen from the available material which comes from Ro-
schütz, near Gera, Thuringia. This locality is also not accessi-
ble now. The hypotypes were chosen from a thin sectioned
material of the same age (Zechstein Formation, Lower Zech-
steinkalk; Upper Permian), made from a core material taken on
the Rügen island.
Locus typicus
: Thieschitz near Gera, Thuringia, Germany.
Stratum typicum
: Zechsteinkalk Ca 1 (bedded facies), Zech-
stein Formation, Upper Permian (? Kazanian).
Additional material
: Six thin sections at the Bundesanstalt
für Geowissenschaften und Rohstoffe, Berlin (BGR); Zech-
stein (Upper Permian), bedded facies. Borehole Samtens 101
(1736.86–1739.35 m and 1744.05–1747.7 m), SW-Rügen,
Germany. Seven colonies and an acetate peel at the Naturkun-
demuseum Gera: III/C2319 (neotype), III/C2615, Roschütz
near Gera, Germany; III/C2627, Röpsen near Gera, Germany;
and one un-numbered specimen from Trebnitz near Gera, Ger-
many; Lower Zechsteinkalk, bedded facies.
Diagnosis
: Discoid colonies; autozooecial diaphragms
rare; apertures with well-developed lunaria; 4–5 spines
in lunaria.
Fig. 1. –
Fistulipora radiata
(S
CHAUROTH
, 1854). – A–B:
Discoid colony upon a brachiopod shell, neotype III/C
2319; scale bars 10 mm for A and 5 mm for B. – C: Discoid
colony upon a brachiopod shell, III/C 2615; scale bar 2
mm. – D: Discoid colony upon a brachiopod shell, III/C
2627; scale bar 10 mm. – E: Colony surface with autozo-
oecial apertures, neotype III/C 2319; scale bar 1 mm. – F:
Longitudinal thin section, hypotype (X 12896); scale bar 1
mm. – G: Tangential thin section, hypotype (X 12896);
scale bar 0.5 mm. – H: Aperture, tangential thin section,
hypotype (X 12896); scale bar 0.1 mm. – I: Tangential thin
section, hypotype (X 12897); scale bar 0.5 mm. – J–K:
Tangential peel, neotype III/C 2319; scale bars 1 mm.

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Description
: Massive discoid colonies, 4.0–7.2 mm in
diameter, 0.45–1.30 mm in thickness. Central part of the
discoid colony usually depressed. Apertures circular,
spaced 4–6 in 2 mm and 5 in 1 square mm of the colony
surface, sometimes contacting with each other. Inter-
spaces between apertures 0.07–0.24 mm. Lunaria well-
developed, directed to the colony centre, 0.12–0.16 mm
wide and 0.09–0.19 mm long, having 0.015–0.018 mm
thick walls, containing 4–5 spines. Lunarial spines
0.018–0.024 mm in diameter. Autozooecial diaphragms
rare. Vesicles generally small, polygonal in tangential
section, separating the autozooecia in 1–3, usually in a
single row; in longitudinal section flattened, having
rounded roofs, spaced 11–13 in 1 mm of the longitudinal
section.
Comparison
: The species
Fistulipora radiata
(S
CHAU-
ROTH
, 1854) is very similar to the species
F.
lunaris
R
O-
MANTCHUK
in M
OROZOVA
1970 from the Upper Permi-
an of the Khabarovsk region (Primorje), Russia.
However, M
OROZOVA
(1970) did not report any spines
in lunaria of this species, which are very distinct in the
present material (Figs. 1G, I). Furthermore,
F. lunaris
has incrusting colonies with multilamellar expansions,
and smaller aperture diameters (0.18–0.22 mm vs. 020–
0.33 mm in
F. radiata
). Another similar species,
Fistu-
lipora enodata
G
ORJUNOVA
, 1975 from the Lower Per-
mian of Tajikistan, differs from
F. radiata
(S
CHAU-
ROTH
, 1854) in having more widely spaced autozooecial
apertures (3–4 vs. 4–6 in 2 mm in
F. radiata
).
Occurrence
: Zechstein Formation, Lower Zechsteinkalk,
bedded facies (Upper Permian); Germany.
Discussion
The bryozoan fauna from the Zechstein is apparently
bound only to the first cycle of carbonate sedimentation
(Ca 1). This corresponds to normal marine conditions,
which subsequently became worse because of restricted
influx of water from the North. The ingression of ocean
waters was short-termed (KIERSNOWSKIE et al. 1995)
and led to an introduction of a restricted fauna within the
newly flooded Southern Permian Basin (ZIEGLER 1990).
Following ingressions were seemingly not accompanied
with an introduction of any bryozoans into the Zechstein
Sea.
The occurrence of a cystoporate bryozoan in the
Zechstein fauna is very interesting in palaeobiogeo-
graphical context. Typical cystoporate genera such as
Fistulipora, Eridopora and Cyclotrypa were widely dis-
tributed during Carboniferous and Permian time being
extraordinary successful in many marine assemblages.
Despite their general abundance and diversity represen-
tatives of this bryozoan group seemed to be abandoned
from the Zechstein Sea. However, several specimens of
Fistulipora radiata (VON SCHAUROTH, 1854) were
found at two distant localities, suggesting a relatively
wide distribution and abundance within the Zechstein
Sea. This species is apparently restricted to the bedded
facies of the Lower Zechsteinkalk. No records from reefs
are known. Such a distribution pattern can also be ex-
plained by a taphonomic effect. Cystoporate bryozoans
mainly possess granular walls in their skeletons which
are sensitive to diagenesis. By contrast, fenestrates and
trepostomes predominantly contain stable laminated
walls. Rocks of Zechstein reefs are usually strongly
dolomitized where most of the delicate internal struc-
tures are destroyed. The bedded facies, however, often
contains rocks and fossils of excellent preservation.
In the Upper Permian, more than twenty species be-
longing to the genus Fistulipora are known world-wide.
Many of them exhibit a morphology similar to that of F.
radiata (SCHAUROTH, 1854). F. lunaris ROMANTSCHUK
in MOROZOVA, 1970 from the Upper Permian (Guadalu-
pian) of Russian Primorje is the morphologically most
similar species. Unfortunately, the record and species
determination in this bryozoan group are far from being
perfect. Thus, the question of the precise ancestry of Fis-
tulipora radiata (VON SCHAUROTH, 1854) awaits its so-
lution.
Acknowledgements
D
IETER
W
EYER
, Berlin, is thanked for his help in finding dis-
covering collections of Zechstein bryozoans in Berlin. F
RANK
H
ROUDA
, Leipzig, helped with information on the collection at
the Naturkunde Museum in Gera and provided useful informa-
Tab. 1. Measurements of the species
Fistulipora radiata
(S
CHAUROTH
, 1854). – Abbreviations: N, number of mea-
surements; X, mean; SD, standard deviation; CV, coefficient of variation; MIN, minimal value; MAX, maximal value.
N X SD CV MIN MAX
aperture width 20 0.25 0.035 13.873 0.20 0.33
lunaria width 20 0.14 0.014 9.940 0.12 0.16
lunaria length 20 0.13 0.030 22.848 0.09 0.19
diameter of vesicles 10 0.08 0.021 26.915 0.06 0.11
vesicles per 1 mm 5 11 1.095 9.781 10 13

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tion. P
RISKA
S
CHÄFER
, Kiel, is thanked for critical reading of
the manuscript and helpful comments. I thank also P
AUL
T
AY-
LOR
, London, for improving the language stile of the manu-
script and helpful comments, as well as two anonymous re-
viewers for their comments.
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