Ernst et al. 2015e

Paläontol Z (2015) 89:645–651
DOI 10.1007/s12542-014-0249-x
SHORT COMMUNICATION
Stratigraphic relevance and typification of the Mississippian
fenestrate bryozoan Narynella narynica (Nikiforova, 1933)
Andrej Ernst • Zoya Tolokonnikova
Patrick N. Wyse Jackson
•
Received: 28 May 2014 / Accepted: 19 September 2014 / Published online: 2 October 2014
Ó Paläontologische Gesellschaft 2014
Abstract The fenestrate bryozoan Narynella narynica
(Nikiforova, 1933) shows high potential for stratigraphic
correlation of the Mississippian deposits of Middle Asia.
This species was recorded in the upper Tournaisian to
lower Viséan deposits of Uzbekistan, Kyrgyzstan and at
various localities in Kazakhstan. New material for N.
narynica is described and type material for this species is
formally designated.
Keywords Bryozoa Mississippian Stratigraphy Taxonomy
Kurzfassung Die fenestrate Bryozoe Narynella narynica
(Nikiforova, 1933) zeigt hohes Potential für stratigraphische Korrelation des Mississippiums von Mittelasien. Diese Art wurde in Sedimenten des oberen Tournaisiums und
unteren Visèums von Usbekistan, Kirgisistan und an verschiedenen Lokalitäten in Kasachstan gefunden. Neues
Material von N. narynica wird beschrieben und Typusmaterial formal festgelegt.
A. Ernst (&)
Institut für Geologie, Universität Hamburg, Bundesstr. 55,
20146 Hamburg, Germany
e-mail: [email protected]
A. Ernst
Tomsk State University, Lenin Ave. 36, 634050 Tomsk, Russia
Z. Tolokonnikova
Kuban State University, 353235Aphipskij,
Post box 30, Krasnodar, Russia
e-mail: [email protected]
P. N. Wyse Jackson
Department of Geology, Trinity College, Dublin 2, Ireland
e-mail: [email protected]
Schlüsselwörter Bryozoa Mississippium Stratigraphie Taxonomy
Introduction
Bryozoans are a phylum of sessile colonial metazoans with a
fossil record going back at least to the Lower Ordovician. In
the Palaeozoic, bryozoans inhabited various marine environments, often being rock-forming. Their skeletons, mainly
of low magnesium calcite, display a high potential for
preservation being useful for various kinds of palaeontological research (Taylor and Allison 1998; Smith et al. 2006).
However, stratigraphic use of bryozoans is quite restricted.
The general obstacles are complex bryozoan morphology
and the necessity of preparation of thin sections for study of
the Palaeozoic stenolaemate bryozoans, as well as relatively
slow species turnovers and restricted distribution. Nevertheless, Bancroft (1987) stressed the importance of Carboniferous bryozoans for biostratigraphy. Especially in the
Mississippian, bryozoans were abundant and diverse and are
well known from various areas worldwide. They contain also
taxa that can be useful for quite fine-scaled biostratigraphic
correlation of distant regions.
The species Fenestella narynica was originally described
by Nikiforova (1933) from the Mississippian of Uzbekistan.
Later, this species was repeatedly reported from the Lower
Viséan deposits of the Middle Asia (Kazakhstan, Kyrgyzstan, and Uzbekistan) (Nikiforova 1933, 1948, 1950; Nekhoroshev 1948; Balakin 1975). Morozova (2001) erected a
new genus Narynella with F. narynica as the type species.
This genus has uniquely shaped autozooecia, which can be
only compared with those of the likewise exotic genus
Lunofenestella Morozova, 2001, from the Viséan of Middle
Asia and Japan (see ‘‘Systematic palaeontology’’).
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The material used for the present study was collected
during the geological mapping in Central Kazakhstan
performed by LTD ‘‘Tsentrgeolsjemka’’ in 2004, by F.
B. Dolgan, senior geologist of the Abai Geological Team.
The bryozoan collection was obtained in the Karagandy
district of Central Kazakhstan, northeast of the Kulanutpes
depression (49°410 1400 , 71°300 0000 ; Fig. 1a), and sent for
identification to the Siberian Palaeontological Centre of
Tomsk State University, Russia. The samples P-4/18 and
P-4/21 were collected from the argillite-carbonate
sequence of late Tournaisian age (State Geological Map of
Fig. 1 a Map showing the
position of the locality where
the studied material was
collected (asterisk).
b Stratigraphic column showing
the position of samples (P-4/18
and P-4/21, asterisks) bearing
Narynella narynica
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A. Ernst et al.
USSR, 1959, 1:500000, M-42-XXIV) (Fig. 1b). Studied
material is deposited at the Laboratory of Micropalaeontology of Tomsk State University, Russia.
Mississippian sediments in Central Kazakhstan are
represented by shallow water organogenic and pelitomorphic limestones, silicified limestones and marls of Tournaisian and early Viséan age. The thickness of these
deposits reaches 0.5 km here, and increases up to 2 km in
the Sarysu-Tengiz depression. This mainly marine
sequence is overlain by carbonate-terrigenous deposits of
the upper Viséan and Serpukhovian (Polyanin and
Stratigraphic relevance and typifications of the Mississippian fenestrate bryozoan
Loginova 2013). The lithological succession was deposited
during the closure of the Palaeozoic oceans and formation
of the north Eurasia landmass (Karyaev 1984). In the
Mississippian, the Tadzhik and Karakum-Tarim continents
were isolated by the Gissar oceanic basin, including the
Gissar island arc. They sequentially approached the early/
middle Palaeozoic continental margin of Kazakhstan and
North Tien Shan. By Moscovian time, the Karakum-Tarim
continent had collided with the Kazakhstan-North Tien
Shan continent, and the Tadzhik continent collided with the
Karakum-Tarim continent (Zonenshain et al. 1990).
The purpose of the present communication is to provide
an updated generic diagnosis of Narynella Morozova, 2001
and to describe new material and discuss the fenestrate
species N. narynica (Nikiforova, 1933) from the Mississippian of Middle Asia, which has a potential to be a
reliable marker fossil for the Mississippian deposits in
Middle Asia.
Systematic palaeontology
Phylum Bryozoa Ehrenberg, 1831
Class Stenolaemata Borg, 1926
Order Fenestrata Elias and Condra, 1957
Family Fenestellidae King, 1849
Genus Narynella Morozova, 2001
Fenestella Lonsdale, 1839 (pars)—Nikiforova 1933, p. 14;
1948, p. 15; 1950, p. 106; 1948, p. 17; Schulga-Nesterenko
1952, p. 29; Balakin 1975, p. 75.
Narynella Morozova 2001, p. 58.
Type species: F. narynica Nikiforova, 1933. Mississippian,
lower Viséan; Naryn Tau mountain ridge, Kyrgyzstan.
Diagnosis: Fan-shaped reticular colonies produced by linear to slightly sinuous, essentially parallel, intermediate to
widely spaced branches joined by narrow dissepiments that
are irregularly spaced. Autozooecia budded in single linear
row but alternately diverging toward frontal surface to
produce two rows per obverse side of branch; low axial
keel present with single row of robust, moderately distantly
spaced nodes. Autozooecial chamber size intermediate;
axial wall absent; single row of chambers in endozone,
circular to subquadrate at reverse wall in tangential sections through deep endozone, transversely elongate ovoid
to subrectangular at mid-chamber depth and may become
subtriangular with long axis transverse at shallow chamber
depth; hemisepta and diaphragms absent. Distal tube short.
Autozooecial apertures circular, partial peristome, and
terminal diaphragms may seal older autozooecia. Extrazooecial skeleton laminated, traversed by abundant, smallto moderate-sized styles; spines or rods of granular core
647
surrounded by laminar sheath extending from reverse side
of branches common. Polymorphs unknown.
Comparison: Narynella Morozova, 2001, differs from
Lunofenestella Morozova, 2001, by having trapezoidal
outline of autozooecia in tangential section instead of halfmoon shaped autozooecia in Lunofenestella, and by the
structure of the keel with keel nodes in one row instead of
two rows of alternating nods in Lunofenestella.
Generic composition: As far as we can tell, Narynella is
monospecific containing only the type species. Morozova
(2001, p. 58) originally included two other species in the
genus: Fenestella leptothecata Schulga-Nesterenko, 1952
and Fenestella shumovi Schulga-Nesterenko, 1952 but this
generic assignment is not accepted here on morphological
grounds. Ken McKinney (unpublished data) on the basis of
a literature assessment considered that the two Permian
taxa Fenestella variabilis (Schulga-Nesterenko, 1952) and
Fenestella unica (Schulga-Nesterenko, 1952) belonged to
Narynella. However, until the original type specimens of
these are examined and their attribution confirmed or
otherwise, it is prudent that they are excluded.
Occurrence: Mississippian (upper Tournaisian–lower Viséan); Kyrgyzstan, Kazakhstan, Uzbekistan.
N. narynica (Nikiforova, 1933); Fig. 2a–d
F. narynica Nikiforova 1933, p. 22, pl. 7, Figs. 2–4; 1948,
p. 27, pl. 8, Fig. 2, pl, 13, Fig. 2; 1950, p. 119, pl. 8,
Fig. 6–6b; Balakin 1975, p. 93–94, pl. 30, Fig. 2.
Fenestella aff. narynica Nikiforova 1933, p. 22, pl. 7,
Fig. 1; Nekhoroshev 1948, p. 30; Nikiforova 1950,
p. 120–121, pl. 8, Fig. 7.
N. narynica (Nikiforova, 1933)—Morozova 2001, p. 58, pl.
16, Figs. 3–4.
Type material: Not formally designated in original publication of Nikiforova (1933). There she illustrated two
specimens in thin-section (CNIGR 184 (pl. 7, Fig. 4) and
CNIGR 185 (pl. 7, Fig. 3) as F. narynica and these constitute the type suite. CNIGR 186 (pl. 7, Fig. 1) was
identified as Fenestella aff. narynica but is not part of the
type suite. Her illustration of the reverse surface of a colony fragment (pl. 7, Fig. 2) is the single specimen from
which the two thin-sections were prepared. Morozova
(1981) indicated that CNIGR 184 was holotype she did not
formally designate it as such. Romanovskaya and Kadlets
(1985, p. 194) in a Catalogue of holotypes in the CNIGR
Museum, St Petersburg, list the specimens figured by
Nikiforova (1933, pl. 7, Figs. 3 and 4) as being the holotype, under the number 93a, b/3341 (=specimens 184 and
185). Under Article 74.6 (ICZN 1999) their action is
considered to be a valid fixation of lectotype by inference.
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Stratigraphic relevance and typifications of the Mississippian fenestrate bryozoan
b Fig. 2 Narynella narynica (Nikiforova, 1933), Mississippian, lower
Viséan, Chatkal’sk Range, Tyan-Shan Mountains, Uzbekistan, PIN
1132/30 (figured by Morozova (2001), pl. 16, Fig. 4). a Tangential
section through endozone and reverse exozone. N. narynica (Nikiforova, 1933), Mississippian, upper Tournaisian, Central Kazakhstan,
P-4/21(3). b–d View of the weathered colony showing branches with
casts of autozooecial chambers
Lectotype: CNIGR 184 and 185, Kshikaindinsk Suite,
lower Viséan, Mississippian; watershed between western
Boronda and Teke-Sekerik, Silbili mountain, Naryn Tau,
Kyrgyzstan.
Remark: The geographical record of the type locality for N.
narynica is slightly perplexing. At the time when the
species was first described, the study area was called
‘‘Turkestan’’, which is a historical name for the territory of
the Central Asia. Nikiforova (1933, p. 22) mentioned the
mountain ridge Naryn Tau (also: Naryn Too) as the geographical area where the locality was situated. This ridge is
part of the Inner Tien Shan, stretches for 130 km, and lies
in Kyrgyzstan along the left bank of the River Naryn (Great
Soviet Encyclopedia, http://bse.sci-lib.com/). The information about the rivers Boronda and Teke-Sekerik was
unavailable. Balakin (1975, p. 93) mentioned this locality
as situated in Southern Kazakhstan, whereas Morozova
(2001, p. 58) wrote that the type locality should be in
Uzbekistan, but in the caption to Fig. 3 of plate 16 she
mentioned Turkmenistan as the geographical locality for
the specimen she indicated as the holotype CNIGR 184
(Morozova 2001, p. 103). The latter is an apparent typing
error, and can be ignored.
Material: Two fragments of reticulate colony, P 4/18(9)
and P 4/21(3). In addition to these specimens, the generic
diagnosis given above is based on the type specimens as
illustrated in Nikiforova (1933), and also on specimen PIN
1132/30 from Chatkal’sk Range, Tyan-Shan Mountains,
Uzbekistan (Fig. 2a).
Description: Reticulate colony formed by straight,
0.3–0.4 mm wide branches joined by 0.20–0.25 mm wide
dissepiments. Fenestrules rectangular, 0.7–0.9 mm wide
and 2.4–2.9 mm long. Autozooecial apertures circular,
0.072–0.095 mm in diameter, spaced 0.15–0.17 mm from
centre to centre. Autozooecial chamber transversely elongate ovoid to subrectangular at mid-chamber depth,
0.13–0.18 mm long and 0.25–0.33 mm wide. Hemisepta
and diaphragms absent. Nodes not observed in the present
material.
Comparison: This material fits very well with the type
material from Uzbekistan showing only minor differences
(branch width 0.30–0.40 mm vs. 0.38–0.44 mm; dissepiment width 0.17–0.22 mm vs. 0.20–0.25 mm; fenestrule
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width 0.70–0.90 mm vs. 0.50–0.56 mm; and fenestrule
length 2.4–2.5 mm vs. 1.7–2.2 mm in the type material).
Occurrence: The present material comes from the Mississippian (upper Tournaisian) rocks of the Central Kazakhstan. The species N. narynica (Nikiforova, 1933) was
previously recorded from the Mississippian of Middle
Asia. These records include the following occurrences.
Lower Viséan: watershed between western Boronda and
Teke-Sekerik, Silbili mountain, Naryn Tau, Kyrgyzstan
(Nikiforova 1933). Upper Tournaisian: west of Chakpaktas
spring, Central Kazakhstan, Balkhash area (Nekhoroshev
1948). Lower Viséan: Baidjansai, Terekty, Karatau,
Southern Kazakhstan (Nikiforova 1948). Lower Viséan:
Mashat River, Talas Alatau, Kazakhstan (Nikiforova
1950). Lower Viséan: right bank of the spring Aksarsai,
left bank of the spring Ispaisai, Uzbekistan (Balakin 1975).
Discussion
Bryozoans are common fossils in Palaeozoic marine
deposits worldwide. Skeletons of Palaeozoic stenolaemate
bryozoans consist of low magnesium calcite, which is
largely resistant against dissolution and diagenesis (Smith
et al. 2006). Therefore, their use for stratigraphy is facilitated. However, bryozoans represent a complex animal
group, which demand thorough study and exhaustive
preparation. Due to the inclination of identifying of Palaeozoic bryozoans on external shape only, many taxa with
uncertain taxonomic assignment were produced previously.
Such an approach decreased the importance of bryozoans
for stratigraphy and palaeobiogeography.
Carboniferous bryozoans have been studied extensively by many authors (e.g., Ulrich 1890; SchulgaNesterenko 1951, 1955; Nekhoroshev 1953; Trizna
1958). Fenestrates in particular, as the most abundant
and diverse bryozoan group in the Carboniferous, hold a
significant potential for stratigraphic correlation. This
group experienced a rapid diversification in the mid-late
Palaeozoic and become widespread in marine biotopes at
this time. Fenestrates are easily recognized in the field
given their distinctive external characters, but such
characters should not be relied on for species discrimination. Morozova (1974, 2001) and Morozova and Lisitsyn (1996) published a series of important revisions of
fenestrates and stressed the importance of internal morphology such as autozooecial chamber shape and presence and shape of heterozooecia. Snyder (1991a, b),
Hageman (1991a, b), and Hageman and McKinney
(2010) contributed significantly to the improvement of
fenestrate taxonomy and delivered reliable tools for
discrimination of fenestrate taxa.
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The species N. narynica (Nikiforova, 1933) has been
repeatedly recorded from the Mississippian of Uzbekistan,
Kyrgyzstan, and Kazakhstan (Nikiforova 1933, 1948,
1950; Nekhoroshev 1948; Balakin 1975). According to
Nikiforova (quoted by Nekhoroshev 1948, p. 30) this
species is characteristic for the upper Tournaisian to the
lower Viséan, whereas Morozova (2001, p. 58) mentioned
only the lower Viséan as the stratigraphic range for this
species. The material studied here comes from the upper
Tournaisian of the central Kazakhstan and supports the
stratigraphic range of N. narynica as upper Tournaisian to
lower Viséan. Furthermore, Morozova (2001) mentioned
Kyrgyzstan as the geographical distribution of N. narynica;
however, no such record was ever published. It is possible,
that this record comes from the unpublished material at the
Palaeontological Institute of the Russian Academy of Sciences in Moscow.
Morozova (2001) established the genus Narynella on the
basis of the unique shape of autozooecia, which makes it
easily recognizable among other fenestrates. The shape of
autozooecial chambers is a crucial character for the discrimination of fenestrate genera (e.g., Morozova 1974).
This shape (usually identified in the mid-tangential section)
can vary broadly—from rectangular, triangular, or pentagonal to pisiform or worm-like. The shape and size of
autozooecial chambers defines the internal morphology of
bryozoans, and so Snyder (1991a) suggested complementing the taxonomic description of fenestrate taxa by
three-dimensional reconstructions of their autozooecial
chambers. Such reconstructions demand certain skills, and
it appears sufficient to record the two-dimensional shape of
autozooecia at various levels through the branch thickness.
A set of external and internal characters of fenestrate colonies must be considered for their taxonomic description,
in order to perform reliable identifications (Snyder 1991a,
b; Hageman 1991a, b).
The unique autozooecial shape of Narynella allows for
their arrangement in one row on the branch. Normally,
autozooecia are arranged in ‘‘Fenestella’’ type bryozoans
in two rows on the branch, side by side or alternating,
which makes them efficient for filtering using the fenestrules as operational subunits (Cowen and Rider 1972). In
most fenestrate taxa, everted lophophores are positioned so
that they occupy space above a particular fenestrule and the
water currents that each generates is not adversely affected
by the action of adjacent lophophores on the other side of
the branch (often separated by a central carina or keel) that
lie above a different fenestrule. Depending on the dimensions of the fenestrules, between four and twenty zooecia
occupy each, and, as such, they operate as a subunit with in
essence the production of one exhalent current passing
from the reverse surface. Despite such a uniserial
arrangement of autozooecia on branches, the apertures of
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Narynella are actually positioned in two alternating rows,
opening on opposite sides of the branch. The dimensions of
autozooecia, their apertures, and their spacing are not significantly different from the majority of related genera
(e.g., Rectifenestella, Spinofenestella, Lunofenestella), so
they have a comparable lophophore size as those genera.
The evolutionary advantage of such a chamber shape is
unknown.
This investigation has demonstrated that bryozoans can
be successfully used for the stratigraphical correlation of
distant regions even if the material is fragmentary and
weathered. The necessary requirement is a detailed
knowledge of bryozoan taxonomy combined with the exact
stratigraphical positions of the referred taxa. However,
knowledge of bryozoan distributions still needs considerable refinement and revision. Numerous publications containing unsatisfactory descriptions reduce the effectiveness
of utilising bryozoans for stratigraphy and palaeobiogeography. To overcome this problem, comprehensive and
precise taxonomic descriptions must be provided so that
subsequent authors can easily recognise collected material.
Acknowledgments LTD ‘‘Tsentrgeolsjemka’’ (Karagandy, Kazakhstan) is thanked for providing samples for the present study. We
thank Paul D. Taylor, London, and Hans Arne Nakrem, Oslo, for their
helpful and constructive reviews. Andrej Ernst thanks the Deutsche
Forschungsgemeinschaft (DFG) for financial support (project ER
278/6.1). This study is a contribution to the IGCP 596 ‘‘Climate
change and biodiversity patterns in the Mid-Paleozoic’’.
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