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Paläontologische Zeitschrift
Scientific Contributions to
Palaeontology
ISSN 0031-0220
Volume 87
Number 3
Paläontol Z (2013) 87:345-395
DOI 10.1007/s12542-012-0161-1
Taxonomic study of spongy spumellarian
Radiolaria with three and four coplanar
spines or arms from the middle Carnian
(Late Triassic) of the Köseyahya nappe
(Elbistan, SE Turkey) and other Triassic
localities
Paulian Dumitrica, Ugur Kagan Tekin &
Yavuz Bedi
1 23
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RESEARCH PAPER
Taxonomic study of spongy spumellarian Radiolaria with three
and four coplanar spines or arms from the middle Carnian
(Late Triassic) of the Ko
¨seyahya nappe (Elbistan, SE Turkey)
and other Triassic localities
Paulian Dumitrica •Ugur Kagan Tekin •
Yavuz Bedi
Received: 20 December 2011 / Accepted: 26 November 2012 / Published online: 12 February 2013
ÓSpringer-Verlag Berlin Heidelberg 2013
Abstract The present article is a taxonomic study of all
spongy spumellarian radiolarian taxa with three and four
coplanar spines or spongy arms occurring in the middle
Carnian from the Ko
¨seyahya section, near the town of
Elbistan, SE Turkey. This fauna is characteristic of the
Tetraporobrachia haeckeli radiolarian Zone, and comes
from an 8 m thick succession of clayey-cherty limestones
occurring at the lower part of the section. In addition, a few
species from the Middle and Upper Triassic from other
areas have been also included in this study to improve
some generic diagnoses, and to better comprehend the
diversity and evolutionary trends of some genera, sub-
families and families. The taxonomy at the generic and
suprageneric levels is based primarily on the types of
microsphere. This new approach allowed new taxonomic
arrangements of genera and suprageneric units, and sug-
gested new phylogenetic relationships among these radi-
olarians and between them and younger radiolarians. The
authors discuss and describe 69 species, of which 37 are
new, and 14 genera, of which three are new (Paraparo-
naella, Pseudangulobracchia, and Ropanaella). The genus
Triassoastrum and others are reinterpreted. All genera
studied are assigned to five subfamilies, of which two are
new (Tetrapaurinellinae and Triassocrucellinae), and two
families (Tritrabidae and Veghicycliidae). Nine species in
open nomenclature are also illustrated.
Keywords Radiolaria Taxonomy Triassic Turkey
Oman Italy Romania Alaska
Kurzfassung Vorliegender Artikel ist eine taxonomische
Studie aller spongio
¨sen Radiolarienarten mit drei und vier
co-planaren Stacheln oder spongio
¨sen Armen, die im mit-
telkarnischen Ko
¨seyahya Profil nahe der Stadt Elbistan,
(SE Tu
¨rkei vorkommen). Die Fauna ist charakteristisch fu
¨r
die Tetraporobrachia haeckeli Radiolarien-Zone und
stammt aus einer 8 m ma
¨chtigen Abfolge mergelig-kiese-
liger Kalke, aus dem unteren Teil des Profils. Zusa
¨tzlich
werden einige Arten der mittleren und oberen Trias von
anderen Lokalita
¨ten beru
¨cksichtigt, um einige Diagnosen
zu verbessern und die Diversita
¨t und evolutiona
¨ren Trends
einiger Gattungen, Unterfamilien und Familien besser zu
verstehen. Die Taxonomie auf dem Niveau der Gattungen
und daru
¨ber basiert in erster Linie auf den Mikrospha
¨ren-
Typen. Dieser neue Ansatz erlaubt neue taxonomische
Zusammenstellungen der generischen und suprageneris-
chen Einheiten und ermo
¨glicht neue und unerwartete
phylogenetische Beziehungen zwischen diesen Radiolarien
sowie zwischen diesen und ju
¨ngeren Radiolarien. Die
Autoren diskutieren und beschreiben 69 Arten (davon 37
neue) und 14 Gattungen (davon 3 neue: Paraparonella,
Pseudangulobracchia, and Ropanella). Die Gattung
Triassoastrum wird neu interpretiert. Alle bearbeiteten
Gattungen ko
¨nnen 5 Unterfamilien (davon 2 neue:
P. Dumitrica
Institute of Earth Sciences, Universite
´de Lausanne, Ge
´opolis,
1015 Lausanne, Switzerland
e-mail: Paulian.Dumitrica@unil.ch
P. Dumitrica
Dennigkofenweg 33, 3073 Gu
¨mligen, Switzerland
U. K. Tekin (&)
Department of Geological Engineering, Hacettepe University,
06800 Beytepe, Ankara, Turkey
e-mail: uktekin@hacettepe.edu.tr
Y. Bedi
Department of Geological Research, 06520 Balgat,
Ankara, Turkey
e-mail: y.bedi@mta.gov.tr
123
Pala
¨ontol Z (2013) 87:345–395
DOI 10.1007/s12542-012-0161-1
Author's personal copy
Tetrapaurinellinae and Triassocrucellinae) und 2 Familien
zugeordnet werden. Neun Arten in offener Nomenklatur
werden ebenfalls illustriert.
Schu
¨sselwo
¨rter Radiolarien Taxonomie Trias
Tu
¨rkei Oman Italy Romania Alaska
Introduction
The very well preserved and very diverse fauna occurring
in the middle Carnian section of the Ko
¨seyahya Nappe
from the eastern Taurides, SE Turkey was, until present,
described in three articles (Tekin and Bedi 2007a,b;
Dumitrica et al. 2010). In the first two articles, the authors
described most of the nassellarian taxa of this fauna, and in
the third they described the Eptingiidae, the Spongosa-
turnaloididae and the Saturnalidae. In the present article, in
order to make known the whole fauna, we describe all the
genera and species of flat spumellarians with spongy test
bearing three and four coplanar spines or arms. In order to
better comprehend the diversity and evolution of these
radiolarians and the ancientry of some internal structures
occurring in the species from the Ko
¨seyahya section, we
include in this article some Triassic species from the latest
Anisian, Ladinian, Carnian, and early Norian from the
Vicentinian Alps (north Italy), Eastern Carpathians
(Romania), Oman, and Alaska.
As already mentioned (Dumitrica et al. 2010), the
importance of the middle Carnian fauna from the
Ko
¨seyahya Nappe lies not only in its diversity, but espe-
cially in its excellent preservation, which allows knowing
the initial skeletal structures of many species that we
consider as primordial, for the building of a much more
natural taxonomic system for the Triassic Spumellaria and
Entactinaria.
Geological setting and material
The main part of the radiolarians presented in this study
comes from the Ko
¨seyahya section, near the town of Elb-
istan, SE Turkey. The section was described and illustrated
in detail in the previous articles (Bedi et al. 2005; Tekin
and Bedi 2007a,b; Dumitrica et al. 2010). It is situated
1 km south of the Ko
¨seyahya village (L38 C2 quadrangle
sheet, between 42.28.360 N/3.62.755 E and 42.28.484 N/
3.62.872 E UTM coordinates), and was measured along
one of the slices of the Ko
¨seyahya nappe. Although the
total thickness of the section is about 118 m and 26 sam-
ples were taken along it, well-preserved radiolarians were
only recovered from five samples (04-ELB-1, 04-ELB-2,
04-ELB-3, 04-ELB-4 and 04-ELB-5). These productive
samples come from unit 2, consisting of an 8 m thick
succession of cherty and clayey limestones overlying unit
1, represented by an alternation of sandstones and marls
(Tekin and Bedi 2007a,b; Dumitrica et al. 2010).
Age of the Ko
¨seyahya radiolarian fauna
Besides nassellarians, entactinarians and spumellarians,
part of which have already been published (Tekin and Bedi
2007a,b; Dumitrica et al. 2010), the productive samples of
the Ko
¨seyahya section contain many characteristic spu-
mellarian and entactinarian taxa of the Middle Carnian
Tetraporobrachia haeckeli Zone, such as: Tetraporobra-
chia haeckeli Kozur & Mostler, Divatella austriaca Kozur
& Mostler, Weverella tetrabrachiata Kozur & Mostler,
Palaeosaturnalis hugluensis Tekin, Spongotortilispinus
carnicus (Kozur & Mostler), Heliosaturnalis magnus
Kozur & Mostler, many species of Veghicyclia Kozur &
Mostler, Capnuchosphaera De Wever, Praeorbiculiform-
ella Kozur & Mostler, Spongosaturnaloides Kozur &
Mostler, Ploechingerella Kozur & Mostler, and many
others as already mentioned, and as we shall show in this
paper and in some forthcoming articles (in preparation).
Compared with the radiolarian faunas of the same zone
from Go
¨stling and Großreifling in Austria (Kozur and
Mostler 1972,1978,1979,1981; Lahm 1984) and the
Huglu unit in Turkey (Tekin 1999), this association is
much richer and much better preserved. It contains many
new species and even genera that were not yet reported
from other areas. This fact is, most probably, a conse-
quence of the very good preservation and of the very
favourable environmental conditions for the development
of radiolarians.
Without waiting for the completion of the study of the
whole fauna; that is, without knowing its complete content,
but based only on the new nassellarians described by Tekin
& Bedi (2007b) and especially (we cite) on the ‘‘co-
occurrence of the species of Elbistanium Tekin, with spe-
cies characteristic of the Tetraporobrachia haeckeli Zone
along with several other taxa not present below this zone
including Castrum spp., Syringocapsa firma Tekin, S. nuda
Tekin, Podobursa spp., Spinocapnuchosphaera spp., and
the first Unumidae’’, Kozur et al. (2009) defined a new
biostratigraphic zone for this assemblage, which they
called the Elbistanium gracile Zone. According to them,
the zone is only known in the Ko
¨seyahya section studied in
the present paper. As the lower boundary of the zone, they
considered the first appearance datum (FAD) of Elbista-
nium gracile Tekin and Goestlingella tueysuezi Tekin.
To this proposal, we make the following remarks:
As in many papers, Kozur, alone or in collaboration,
described new radiolarian zones not based on more or less
continuous sections, where one can see the succession of
346 P. Dumitrica et al.
123
Author's personal copy
species and the FADs or last appearance datum (LAD)s of
marker species, but from small, disparate sections, or only
from single samples with rich assemblages. It is not only
thecaseoftheE. gracile Zone, but also of the Spongo-
tortilispinus moixi Zone (Moix et al. 2007)andmany
others. As a result, the FADs or LADs he proposed do not
represent the true FADs or LADs of the respective spe-
cies, but probably their acme or just occurrences. For
instance, we do not know yet if the FADs of E. gracile
and G. tueysuezi are synchronous as Kozur et al. (2009)
wrote. What we do know is that the two species co-occur
in the Ko
¨seyahya section samples. Also, we do not know
when either species first occurred. For instance, the genus
Elbistanium Tekin was described on the basis of the
Elbistan fauna and was thought to have first appeared at
this level, but the senior coauthor has already seen a
specimen of this genus in the well-known early late
Longobardian sample 88/272 from Fojnica (Bosnia) sev-
eral years ago in Prof. Mostler’s collection, at the Uni-
versity of Innsbruck. So the FAD of this genus is not late
Julian, but late Ladinian (early late Longobardian), when
many Carnian genera have their FADs. The same thing
can be said regarding the FAD of Goestlingella tueysuezi.
It is very probable that its FAD is much earlier than that
we know today from the Elbistan samples, because the
youngest species of this genus that we knew before the
publication of this species (Goestlingella goricanae Tekin
& Mostler and G. pseudoillyrica Tekin & Mostler) were
also early late Longobardian, based on the very rich
sample from Fojnica. We know no other species of this
genus in the latest Longobardian and Cordevolian (early
Julian), because of the less good preservation of the
radiolarian faunas occurring in this interval. It is possible
that G. tueysuezi appeared earlier, or that the upper range
of the Longobardian species also covered the lower Julian
to give rise to this late Julian species. It is also possible
that other species, unknown until present, made the con-
nection between the late Longobardian and the late Julian
species.
The upper boundary of the E. gracile Zone was con-
sidered by Kozur et al. (2009) to correspond with the FAD
of S. moixi Kozur & Mostler, Karnospongella multispinosa
Kozur, Moix & Mostler, Zhamojdasphaera rigoi Kozur,
Moix & Mostler, and also the FAD of more than 100
species described from the very rich early Tuvalian (early
Late Carnian) sample G11 collected by Patrice Moix and
partly studied by Moix et al. (2007) and Kozur et al.
(2007a,b,c,2009). The synchronous FADs of more than
100 species are impossible in a normal sequence. It can
only be recorded when:
a) there is a relatively long stratigraphical gap between
two superposed faunas;
b) the samples below a very rich sample contain poorly
preserved faunas;
c) the radiolarian fauna from a lower sample, even if
very rich and very well preserved, is only partly
studied.
The explanation of so many synchronous FADs in the
sample G11, the fauna of which has been studied by Kozur
and coauthors (l. cit.), can be found in almost all the
mentioned cases as follows:
a) The fauna of the sample G11 does not come from a
level immediately above our productive section (sam-
ples 04-ELB 1–5). Since the G11 sample comes from
another locality, we do not know how long the missing
interval between it and our samples is. What we surely
know is that the G11 fauna is a little younger than the
Ko
¨seyahya fauna. But we do not know how big this ‘‘a
little’’ is. What we can say to suggest the time span of
this interval is that on the interval of 5 m represented
by the radiolarian-bearing Ko
¨seyahya section, there is
practically no notable difference between the assem-
blage of the lowermost sample (04-ELB 1) and that of
the uppermost sample (04-ELB 5). The differences we
could notice between them are especially due to
differences in preservation.
b) Until present, there is no section with a fauna of
similar preservation and richness within the strati-
graphic interval between the two faunas. In Oman, in
this interval, the preservation is rather poor, and
consequently, the radiolarian assemblage is also poor
by comparison with the two faunas under discussion.
c) The Elbistan fauna is only partly studied, and many
species or genera Kozur et al. (2009) mentioned as
having their FADs in the sample G11 also exist in the
Elbistan fauna or even earlier. To give only two
examples: Karnospongella multispinosa Kozur, Moix
& Mostler, considered by them to be one of the three
marker species, the FADs of which mark the upper
boundary of the Elbistanium gracile Zone, also occurs
in the Elbistan assemblage, but was not yet mentioned
until present, because it will be treated in another
paper together with other related species and genera;
and the FAD of the genus Spinomersinella Kozur,
Moix & Mostler is not in the Tuvalian sample G11, but
in the uppermost Anisian or lower Ladinian of the
Buchenstein Formation, where it is represented by a
still undescribed species (see O’Dogherty et al. 2009).
In conclusion, we continue considering this fauna as
belonging to the Tetraporobrachia haeckeli Zone, as
described by Kozur and Mostler (1994: 40, 166), and as
being of late Julian (middle Carnian) age, with the idea of
subdivision of the Carnian into two substages (Julian and
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 347
123
Author's personal copy
Tuvalian) (Ogg 2004). The same age is also supported by
the conodont fauna. According to Dr. Leopold Krystyn
(personal communication), the sample 06-MY-10 from the
top of the grey thin bedded limstone, about 2 m above the
uppermost radiolarian sample (04-ELB 5), (see Tekin &
Bedi 2007 for the position of both samples), contains a
small conodont set of Gladigondolella tethydis, ramiforms
of Gladigondolella multi-elements, and Paragondolella
foliata, which is clearly a pre-Late Carnian fauna. He
considers, therefore, that the Ko
¨seyahya radiolarian fauna
corresponds to the topmost Austriacum Zone.
Additional samples
To better understand the diversity and evolution of the
genera and species discussed in this paper, we include
herein some species from older or younger Triassic levels.
The additional samples studied are as follows:
I. Buchenstein Formation, Vicentinian Alps, Italy
Rc4. Buchenstein Formation, latest Anisian, Ladinocampe
multiperforata Zone, probably Ladinocampe annuloperfo-
rata subzone, about 500 m northeast of Monte Anghebe,
along the road running to Recoaro, Vicentinian Alps,
Southern Alps, North Italy (Epting et al. 1976; Dumitrica
1978,1982).
II. BV 85-70. Livinallongo Formation, Monte de Saline,
Marmolada Massif, North Italy (Kellici & De Wever 1995)
According to these authors, the sample comes from the
upper part of the section from Monte de Saline, which
exposes the so-called ‘‘Livinallongo series’’, represented
by tuffaceous marls. Radiolarian fauna studied by these
authors is either latest Anisian or early Ladinian, and
corresponds to what was considered Fassanian before
establishing the base of the Ladinian at the base of the
E. curionii Ammonoid zone (Brack et al. 2005); radiolarian
zone unknown.
III. Transylvanian Nappe, Rarau Mountain, Rarau
Syncline, Eastern Carpathians, Romania
The Triassic radiolarian species and specimens we used in
the present paper come from the radiolarian-bearing rocks
occurring in the Rarau Mountain, in what Patrulius (1996)
separated as the Zimbru ‘‘Series’’ or Zimbru Unit, which is
part of the Transylvanian Nappe of Uhlig (1907). This unit
consists of a series of klippen whose restored sequence
includes, in ascending order: Daonella jasper beds (Fass-
anian to middle Longobardian), Zimbru Limestone (upper
Longobardian—Julian), Popi Limestone (uppermost Car-
nian to middle Norian), Salinaria Marls (upper Norian),
dark grey to black Limestone (Rhaetian), and others.
Radiolarian occurrences are restricted to the first two
members, from which Dumitrica (1982) described two
oertlispongid faunas.
Sample R78-6. About 100 m east of the base of the big
Triassic limestone olistolith of Piatra Zimbrului (Popescu
and Patrulius 1964). Nodular limestone with Daonella
indica Bittner and a radiolarian assemblage indicative of
the Muelleritortis cochleata Zone, Spongoserrula rarauana
Subzone, which is indicative of the early middle Longo-
bardian (Kozur and Mostler 1996). The sample seems to be
older than the following three samples, because it is also
characterized by the occurrence of wide-winged species of
Pterospongus, whereas the genera Tritortis and Steigeri-
spongus are still missing.
Sample R88. Rarau Syncline, Pojorita village, right side
of the Fundu Pojoritei valley. Red chert on the top of an
ofiolite olistolith. The assemblage contains a rather rich
radiolarian fauna, consisting of Bogdanella praecursor
(Kozur & Mostler), Hungarosaturnalis multispinosus
(Kozur & Mostler), Muelleritortis cochleata (Nakaseko &
Nishimura), M. expansa (Kozur & Mostler), Pseud-
ostylosphaera imperspicua (Bragin), Spongoserrula
rarauana Dumitrica, Scutispongus undulatus (Dumitrica),
S. latus (Kozur & Mostler), Tritortis dispiralis (Bragin),
Tubospongopallium gracile (Tekin & Mostler), and many
others (Dumitrica 1982; Hoeck et al. 2009), indicating
a middle Longobardian age, M. cochleata Zone and
S. rarauana Subzone, respectively.
Sample R106. Curmatura Raraului (Rarau Saddle),
Rarau Mt. Block of dark brown cherty limestone in an
escarpment of megabraccia. The radiolarian assemblage is
rather rich and, besides the oertlispongid species described
by Dumitrica (1982), it includes: Muelleritortis cochleata
(Nakaseko & Nishimura), M. firma (Gorican), Tritortis
dispiralis (Bragin), T. kretaensis (Kozur & Krahl),
Pseudostylosphaera nazarovi (Kozur & Mostler), etc. This
is the youngest radiolarian sample from the Rarau Mt.
because it contains not only T. dispiralis, but also many
specimens of T. kretaensis (Kozur & Krahl). Dumitrica
(1982) considered the age of this sample as Tuvalian, but
he agrees now with Kozur and Mostler (1996) who rejected
this age. The age, according to the cited species is latest
Longobardian-earliest Julian or early Cordevolian, the
T. kretaensis Zone.
Sample R114-4. Wedge of about 10 m of red, green and
black chert interbedded with filamentous limestone, in an
escarpment of megabreccia. Curmatura Raraului (Rarau
saddle). Both the red chert and limestone contain Daonella
pichleri Mojs. The radiolarian assemblage is comparable
348 P. Dumitrica et al.
123
Author's personal copy
with that of the sample R78-6, and the age is also similar to
that of the mentioned sample (Dumitrica 1982).
IV. Zulla Formation, Chert Member, Hamrat Duru Group,
Hawasina Complex, Wadi Bani Khalid section, Oman,
UTM 70.65.11 E/24.96.352 N
According to Cooper (1987), Bernoulli et al. (1990) and
Blechschmidt et al. (2004), this formation comprises four
members and spans most of the Triassic as follows:
Limestone and Shale Member (Olenekian—Anisian),
Sandstone and Shale Member (Anisian), Chert Member
(late Anisian—early Norian), and Halobia Limestone
Member (early—late Norian.
The section was described in Blechschmidt et al. (2004),
and partly in Dumitrica and Hungerbu
¨hler (2007). It offers
the best outcrop of the Radiolarian Chert Member of the
Triassic Zulla Formation. In this section, this member has a
thickness of about 55 m and consists of thin-bedded, red
and green radiolarian ribbon chert with interbedded sili-
ceous shale. Its lower boundary is gradual and its top is
marked by a coquina level with Halobia beyrichi Mojs.
The chert member spans the late Anisian (Illyrian)—early
Norian stratigraphic interval. In this article, we illustrate a
few species from this section, occurring in the following
eight samples: BR910,BR911,BR913,BR914,BR917,
BR918, BR919, and BR921. The samples BR910–BR914
are from the Tetraporobrachia haeckeli Zone; BR917 is
from an unnamed zone between T. haeckeli Zone and
S. moixi zone, and its age is probably early late Carnian
(Dumitrica and Hungerbu
¨hler 2007); BR918 and BR919
are early Late Carnian corresponding to the Spongotortil-
ispinus moixi Zone of Moix et al. (2007); and BR921 is late
Tuvalian (late Carnian) with Poulpus carcharus Sugiyama,
Xiphotecaella rugosa (Bragin), Capnuchosphaera spp.,
etc., but without species of the genus Capnodoce Pessagno.
V. Haliw Formation, Hamadiyin I section, Oman, UTM
58.28.11 E/25.21.071 N
The section is located northwest of the Hamadiyin and
Haliw villages, a few hundred meters north of the road
Izki-Sinew. It exposes, from bottom to top, the two for-
mations of the Umar Group of the Hawasina Complex: the
Sini Formation composed mainly of volcanic rocks and the
Haliw Formation (Glennie et al. 1974) or Aqil Formation
(Be
´chennec et al. 1992). The Haliw Formation is composed
of Triassic-Cretaceous predominantly siliceous pelagic
deposits and calcirudite and/or megabreccia with reworked
shallow-marine carbonates. The Triassic portion exposed
in this section is represented by its two lower members:
lower cherty limestone member (about 13 m) of middle
Carnian age, and filamentous red chert member (about
15 m) with rare levels of filamentous limestone of late
Carnian (Tuvalian)—early Norian age. The sample HA4,
which we used for this paper, was collected from the top of
the lower member, at 13 m above the base of the measured
section, and the sample HA 10 at 26.5 m above the base. It
contains Mostlericyrtium sitepesiformis Tekin, Capnodoce
spp., Kahlerosphaera norica Kozur & Mock, and many
other species suggesting a latest Carnian—earliest Norian
age.
VI. Glenn Shale Formation, east-central Alaska
The Glenn Shale of east-central Alaska is composed of two
distinct temporal stratigraphic entities of significantly dif-
fering origins: a lower portion consisting of a 200 m thick
Middle to Upper Triassic marine carbonate unit recognized
throughout east-central Alaska, and an upper portion,
1,600 m thick, of Lower Cretaceous clastic flysch-like unit
(Brabb 1969). The Triassic Glenn Shale of the area located
between the latitudes of 65°000and 65°300N and longitudes
of 141°000and 142°000W was investigated by B. E. Rob-
inson (1988) for the radiolarians studied in his unpublished
Master of Science thesis of the University of Texas at
Dallas. Portions of a residue of one sample were given by
B. E. Robinson to the participants of the 5th Radiolarian
Symposyum held in Marburg in 1988. The sample of
radiolarians that we studied was from one such portion
given to Dr. Spela Gorican who, in turn, gave it to the
senior coauthor (PD) of the paper. The exact location of the
sample in east-central Alaska is unknown; it is from one of
the two measured sections by Robinson (Trout Creek or
Michigan Creek). Radiolarians are very well preserved
and, according to Robinson, belong to the Latium paucum
subzone of the Capnodoce Zone of Pessagno et al. (1979)
and Blome (1984), ranging from latest Carnian (late
Tuvalian) to earliest Norian (early Lacian).
Taxonomy
Until present, the taxonomy of the radiolarians with spongy
skeleton studied in this article was based on external
skeletal morphology: the shape of skeleton and the number
of projections (spines or arms) and their shape and spatial
arrangement. This is the easiest way to determine species
for biostratigraphic purposes, especially when the preser-
vation is not perfect. However, if one wants to make a more
natural taxonomy at the generic and suprageneric levels (as
we tried to do in the present paper) to understand the
relationships among higher taxa and their evolution, one
needs to know the innermost or initial skeletal structures,
which are the most conservative and, consequently, the
most apt for such reconstructions (Dumitrica 1983,1988,
1989,1991,1997,2001; Dumitrica and Carter 1999,
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 349
123
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Dumitrica in De Wever et al. 2001; Dumitrica and Zu
¨gel
2002; Bjørklund et al. 2008). The knowledge of these
internal or initial skeletons is not possible for most fossil
radiolarians, because of their poor preservation. Even if the
outer skeleton is very well preserved, in most fossil radi-
olarians, the initial skeletons, which usually are very deli-
cate, are destroyed by fossilization. That is why it is
necessary to maximally exploit the possibilities offered by
the very well preserved species, as has been the case for
those of the radiolarian fauna from the Ko
¨seyahya section.
This is what we tried to do in the present paper and what
we shall also try to do in a series of papers in preparation,
based on the study of different groups of spumellarian and
entactinarian Radiolaria of this fauna.
It is possible that the taxonomy at higher levels based on
internal or initial structures will be more difficult to accept
immediately, because we are still accustomed with a tax-
onomy based on external morphology. However, the initial
structures oblige us to put together in the same supragen-
eric units (subfamilies, families or superfamilies) genera
with different types of shells and external projections
(spines or arms, etc.), and to take into account the number
of such projections when they are closely connected with
primary beams. Moreover, they oblige us to taxonomically
separate genera with similar external spines or arms; for
example, to consider that Triassocrucella Kozur & Mostler
is not a hagiastrid as Crucella Pessagno, Pseudocrucella
Baumgartner, Higumastra Baumgartner, or other post-
Triassic hagiastrid genera. For generic and suprageneric
taxonomy, we should always have in mind that homeo-
morphy is very common among radiolarians, and that
external morphology is deceptive in many cases and cannot
be considered as a first order taxonomical character. A
beginning has already been made in De Wever et al. (2001)
for the families Pseudoaulophacidae, Angulobracchiidae,
Tritrabidae, etc. A lot of work still remains to be done,
because the study of internal structures is time consuming
and, as they are usually very delicate, the chance to find
well-preserved internal structures is rather reduced in fossil
materials.
Most taxa we describe in the present paper, although
very different morphologically and very diverse taxonom-
ically, have several elements in common: a more or less
organized spongy skeleton, usually three to four coplanar
spines or arms, and two main types of microspheres. In
order to know the ancientness of these microspheres, their
variation and the skeletal morphologies in which they
occur, we extended our study to some taxa comprised
between the uppermost Anisian (the older Fassanian sub-
stage), Ladinocampe multiperforata Zone, Ladinocampe
annuloperforata Subzone, and the lower Norian (see the
list of additional samples above). The results are surprising,
and have shown how far we still are from understanding
the taxonomy, phylogeny, and evolution of spumellarian
Radiolaria.
All radiolarians studied in the present article are
spumellarians and start their skeleton with a microsphere.
The morphological variety of tests of these radiolarians has
its origin in two types of microspheres, as described below:
tritrabid and veghicycliid. In the present article, each of
these types represents the base on which the taxonomy at
subfamily and family levels is built.
Concerning the taxonomic value of the structural and
morphological characters taken into account in the present
article, we preferred to give a family level value to the
type of microsphere and to the number of primary beams
originated in it; a subfamily value to the type of projec-
tions (spines or spongy arms) aligned with the primary
beams; a generic value to the structure of these projec-
tions (spines circular in cross-section or bladed, simple or
with verticils of spinules, etc.); and a species level to the
external morphology. Consequently, the generic and su-
prageneric taxa of the species described in the present
paper is as follow:
1. Family Tritrabidae Baumgartner: with microsphere of
tritrabid type:
1a. Subfamily Intermediellinae Lahm: with spines
aligned with the primary beams;
Paurinella Kozur & Mostler: with spines circular in
cross-section;
Spinopaurinella Kozur & Mostler: with branched
spines circular in cross-section;
?Neopaurinella Kozur & Mostler: with asymmetric
test and a pylome near one spine;
Zhamojdasphaera Kozur & Mostler: with foliaceous
spines;
Angulopaurinella Kozur & Mostler: with three-bladed
spines;
1b. Subfamily Tritrabinae Baumgartner: with arms
aligned with the primary beams:
Triassoastrum Kozur & Mostler emend.: with arms
lacking spongy cortical test;
Paraparonaella Dumitrica & Tekin, n. gen.: with
thick spongy test around central core;
Pseudangulobracchia Dumitrica & Tekin n. gen.:
with arms rectangular in cross section;
?Ropanaella Dumitrica & Tekin, n. gen.: without
primary beams.
2. Family Veghicycliidae Kozur & Mostler, emend.
herein: with pentagonal dodecahedral microsphere and
four primary beams:
2a. Subfamily Tetrapaurinellinae Dumitrica & Tekin,
n. subfam.: with four spines aligned with primary
beams:
350 P. Dumitrica et al.
123
Author's personal copy
Tetrapaurinella Kozur & Mostler: with conical
spines;
Plafkerium Pessagno: with simple three-bladed
spines;
Tetraspongodiscus Kozur & Mostler: with a verticil of
spinules on three-bladed spines.
2b. Subfamily Veghicycliinae Kozur & Mostler: with
circular test and primary beams included in it:
Veghicyclia Kozur & Mostler: with latticed equatorial
plate and small spongy test around microsphere;
Carinacyclia Kozur & Mostler: with equatorial plate
covered on both sides by latticed or spongy test;
Praeorbiculiformella Kozur & Mostler: with a thick-
ened circular peripheral test and equatorial peripheral
spines.
2c. Subfamily Triassocrucellinae Dumitrica & Tekin n.
subfam.: with four arms aligned with primary beams:
Triassocrucella Kozur: with arms terminated into
three spines, of which one is axial and the other two
are lateral.
Taxonomic descriptions and discussions
All holotypes and paratypes of the new species described in
the present paper are mounted on stubs together with other
unpublished taxa in the collection of the senior coauthor,
and will be stored in the collection of the Muse
´ede
Ge
´ologie Lausanne.
Class Radiolaria Mu
¨ller 1858
Subclass Polycystina Ehrenberg 1838, emend. Riedel 1967
Order Spumellaria Ehrenberg 1875
1. Family TRITRABIDAE Baumgartner 1980, emended herein
Type genus:Tritrabs Baumgartner 1980: 293.
Emended diagnosis: Spongy Spumellaria with three
coplanar spines or spongy arms bearing a small triangular
microsphere with corners lying in the equatorial plane, one
laterally directed interradial pore on each side between
corners and, on each face, with a three-rayed structure or a
little more complicated structure connected to the frames of
the interradial pores.
Remarks: The tritrabid microsphere was described and
illustrated by Dumitrica in De Wever et al. (2001, Fig. 85),
based on the study of Jurassic and Cretaceous species. At
that time, this structure was only known in the three-armed
genera with arms of hagiastrid structure (Tritrabs Baum-
gartner, Archaeotritrabs Steiger, and Neoparonaella Yang).
The occurrence of the same microspheric structure in some
Triassic genera with a spongy shell armed with three spines
allows us to extend the definition of this group, and to see the
origin of the true Tritrabidae in the Middle Triassic genera
with spines instead of hagiastrid-type arms. This micro-
sphere (Fig. 1c) is a very simple triangular structure with
nine pores (three interradial or lateral pores opened in the
interradial area of the initial skeleton, three on one side of
the equatorial plane, and three on the other side, opened on
the direction of the primary beams); a three-rayed structure
on each face connecting the upper and lower frames of the
interradial pores, and a bent bar at each corner of the triangle
lying in the equatorial or subequatorial plane. The deviation
of these bent bars from the equatorial plane is due to the
force exercised by the two adjoining microspheric bars
arising from the equatorial ring, of which one goes up and
the other goes down. These bent bars are at the origin of the
three primary beams (pr) characteristic of the Tritrabidae.
Of all these characters, the presence of the three interradial
pores (one on each side of the triangle) and the bent corners
in the equatorial or subequatorial plane in which the three
primary beams originate are the most important and constant
characters of this type of microsphere. They structurally
differentiate the members of this family from the members
of the family Pseudaulophacidae (Dumitrica 1997;De
Wever et al. 2001, Fig. 83), where there are also three pri-
mary beams but no triangular structure in the equatorial
plane; the primary beams originating in a bent vertical mi-
crospheric bar. Moreover, in the latter family, there is a
triangular ring on each face of the microsphere that may
become hexagonal by the appearance of a bar in the middle
of each side of the triangular ring connecting the micro-
sphere with the first perimicrospheric girdle.
When the tritrabid structure was first illustrated (De
Wever et al. 2001), we knew neither how old this structure
was, nor how common and how diverse it was in the Tri-
assic. The only species on which the Triassic (Norian) FAD
was recorded at that time was Paronaella norica Kozur &
Mock (assigned in the present article to the genus Trias-
soastrum Kozur & Mostler emend. herein) occurring in the
latest Carnian—early Norian sample from the Glenn Shale
Formation of east-central Alaska (Fig. 2a–c). The very
well preserved fauna from the Ko
¨seyahya section in the
Elbistan area provided us the possibility of finding well-
preserved initial skeletons of many species, and conse-
quently, of better understanding the phylogenetical
relationships of many genera. To have a better image of the
occurrence of this microsphere in the Triassic radiolarian
fauna, and to know its ancientness and its variability, we
made additional sections in some older and younger
Triassic species (Fig. 1). As a general rule, we found that in
the Triassic, almost all microspheres of this type are less
regular and less simple than those occurring with the
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 351
123
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Jurassic and Cretaceous species. They have the general
structure of the tritrabid microsphere (three corners in the
equatorial or subequatorial plane and three interradial
pores), but their upper and lower faces are neither as reg-
ular nor as simple as in the Jurassic and Cretaceous species.
Usually, the two faces (upper and lower) of the Triassic
microspheres are unequal: one face may be simple, with a
three-rayed structure as in the Jurassic and Cretaceous
species, whereas the other face can have more bars and,
accordingly, more pores. This fact proves the primitive
character of this type of microsphere in the Triassic, and
suggests that its simplicity and regularity in the Jurassic
and Cretaceous are secondary features resulting from a
process of reduction of some skeletal elements and increase
of symmetry.
Occurrenceof this structure in some Triassic genera with a
spongy shell and three spines or arms allows us to extend the
definition of this group, and to see the origin of the true
Tritrabidae in the Middle Triassic genera with spines instead
of hagiastrid arms. In this situation, we think it necessary to
distinguish in this family two subfamilies considered as
evolutionary stages of the group: the subfamily Intermedi-
ellinae Lahm, including genera with spongy shell and three
spines, and the subfamily Tritrabinae Baumgartner, including
genera with complex arms of hagiastrid structure. In the
evolution of this group, one can note a change of the structure
of the radial projections from robust spines with circular cross
section to bladed spines, and finally to spongy arms devel-
oped around the primary beams. It is an evolution also noted
in the other families treated in this article.
Fig. 1 Microspheres of
tritrabid type of some Middle
and Late Triassic species (a,b,
d–h) showing its variability and
the asymmetry of the two faces
by comparison with the
microspheres of Jurassic and
Cretaceous species (c). a
Paurinella aequispinosa Kozur
& Mostler. bAngulopaurinella
edentata Dumitrica & Tekin n.
sp. cSchematic microsphere of
tritrabid type, redrawn after De
Wever et al. (2001).
d,eTriassoastrum trammeri
(Kozur & Mostler).
fZhamojdasphaera latispinosa
Kozur & Mostler.
gAngulopaurinella
dentispinosa Dumitrica & Tekin
n. sp. hAngulopaurinella
edentata n. sp. Scale bar is for
all figures except c
352 P. Dumitrica et al.
123
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Range: Middle Triassic (middle Anisian) to Late Creta-
ceous (Maastrichtian).
1a. Subfamily INTERMEDIELLINAE Lahm 1984
Syn. Zhamojdasphaeridae Kozur & Mostler 2006, type
genus: Zhamojdasphaera Kozur & Mostler 1979.
Type genus:Intermediella Lahm 1984:53=Paurinella
Kozur & Mostler 1981: 49.
Diagnosis: Tritrabidae with globular to lens-shaped cir-
cular to triangular spongy skeleton armed with three spines
that are either massive or two- or three-bladed. On the two
faces, the structure of the microsphere is either simple as in
Tritrabs or often it may have more bars and pores, but it
always has three laterally opened pores in the interradial
areas, and the bars from the three corners are always in the
equatorial or subequatorial plane.
Remarks: In the present state of knowledge, it is difficult
to know the diversity of this subfamily during the Triassic.
Sections that the senior coauthor made in a few Fassanian
or Illyrian species prove that the family comprises not only
the type genus, but also the genus Zhamojdasphaera Kozur
& Mostler and the genus Angulopaurinella Kozur &
Mostler. Zhamojdasphaera, with its foliaceous or two-
bladed spines, is just a peculiar case and it is not necessary
to erect a special family for it, as we do not erect a special
family for the Oertlispongidae with foliaceous spines to
separate them from those with circular cross-section spines.
Unfortunately, due to the poor preservation of the most
internal skeletons of the specimens we studied, we failed to
see the microspheric structure of the type species of the
genus Intermediella. However, we could see it in some
species of the genus Paurinella, as described by Kozur and
Mostler (1981). The spongy shell of the species of this sub-
family is either concentrically disposed (Fig. 3b, e, g), or more
or less irregular(Fig. 3a, c, d), sometimes also with a tendency
towards a radial disposition (Fig. 3a–c). It seems that the
concentric disposition is characteristic of some genera
(Zhamojdasphaera) whereas in other genera (Paurinella)it
may be either concentric or irregular, depending on species.
Since in this paper, the systematics at the family level is
based on the type of microsphere, we do not follow the
Fig. 2 a–cTriassoastrum
trammeri (Kozur & Mostler),
Glenn Shale, Alaska.
aMicrosphere and proximal
part of primary beams with
position of their successively
inverted verticals of branches.
bTransversal section through
arms showing the reciprocally
inverted position of branches of
first, second and third layer
around the primary beam for
two successive levels of
branches. cPosition and
nomenclature of pores in
transversal section through an
arm: 1primary pores, 2
secondary pores, 3tertiary
pores. Bar only for Fig. 2a.
dTransversal section through
an arm of Pseudangulobracchia
prisca Dumitrica & Tekin n. sp.
eTransversal section through an
arm of Paraparonella okuyucui
Dumitrica & Tekin n. sp
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 353
123
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emended diagnosis made by Kozur and Mostler (1994).
Consequently, we exclude from this subfamily the genera
Katorella Kozur & Mostler, Discokatorella Kozur &
Mostler, and Tetrapaurinella Kozur & Mostler that have
four spines and a microsphere with 12 pores (see below). In
the present article, we include them in the new subfamily
Tetrapaurinellinae of the family Veghicycliidae. Although
we could not make sections in all these four-spined genera
to know their microsphere, the number of spines suggests
that they should have the microsphere of the Tetrapauri-
nella type.
Range: Middle Anisian to Early Norian so far as known.
Genus Paurinella Kozur & Mostler 1981
354 P. Dumitrica et al.
123
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Syn. Intermediella Lahm 1984 (type species: Intermediella
trispinosa Lahm 1984).
Type species:Paurinella curvata Kozur & Mostler 1981.
Remarks: Until present, there was no information on the
structure and morphology of the microsphere. Sections the
senior coauthor made in a specimen of Paurinella aequi-
spinosa Kozur & Mostler have shown that its skeleton
starts from a microsphere of tritrabid-type (Fig. 1a).
Although we do not know if this situation is general, we
have reason to think that the structure of the microsphere is
reflected in the test morphology and, therefore, that all
species of the genus Paurinella with equal and equally
spaced spines have the same type of microsphere.
Range: Middle Anisian to late Carnian.
Paurinella aequispinosa Kozur & Mostler 1981
(Figs. 1a, 3b)
1981 Paurinella aequispinosa Kozur & Mostler: 50, pl.
42 Fig. 1, pl. 43 Fig. 1
1994 Paurinella aequispinosa Kozur & Mostler.—Kozur
& Mostler: 72, pl. 15 Figs. 9, 11
Remarks: The microsphere of this species is heteropolar
in polar direction, one face having the three-rayed structure
as in the post-Triassic Tritrabidae, whereas the other face
has three pores separated in the central part by a three-
rayed structure, inverted by comparison with the same
structure from the opposite face. The interradial pores are
always present. The spongy test is disposed in five or more
concentric layers, resembling that of Zhamojdasphaera
latispinosa Kozur & Mostler 1979 (Fig. 3g).
Range and occurrence: Latest Anisian to early Ladi-
nian, Southern Alps, N Italy.
Paurinella curvata Kozur & Mostler 1981
(Fig. 4a)
1981 Paurinella curvata Kozur & Mostler: 49, pl. 4
Figs. 3,6
Remarks: A section that we made in this species gave no
positive result, because the centre of the shell was partly
destroyed during the preparation of the section. Although
the spongy test is rather massive but with a certain radial
disposition, along the spines one can see that it is organized
in up to 14 layers forming a corresponding number of
circumferential rings.
Range and occurrence: Latest Anisian to early Ladinian
(Fassanian), Buchenstein Formation, Recoaro, Vicentinian
Alps, N. Italy.
Paurinella mutihaci Dumitrica & Tekin n. sp.
(Fig. 4b)
Description: Test spongy, lens-shaped, inflated, circular
in face view. Test surface with numerous small pores.
Spines cylindrical on the proximal part and conical on the
distal one, with a gradual transition between the two
portions.
Material: Two specimens, of which only the holotype
was photographed.
Holotype: Fig. 4b, sample R78-6, Rarau Mt., Eastern
Carpathians, Romania, Muse
´edeGe
´ologie Lausanne, No.
MGL 96814.
Dimensions (in lm, based on holotype): Diameter of
spongy skeleton 145, length of spines 120, diameter of
spines 36.
Remarks:Paurinella mutihaci n. sp. differs from P. tri-
angularis Kozur & Mostler in having a circular rather than
a triangular shape, and in having a cylindrical proximal
portion of spines, whereas the spines of the latter species
increase in diameter up to the middle portion where the
maximum diameter can be measured. It also differs from
P. aequispinosa in having spines that are shorter, thicker
and that are not gradually tapering.
Etymology: The species is named for the Romanian
geologist Vasile Mutihac, to honour his contribution to the
geology of the Rarau syncline, Eastearn Carpathians.
Fig. 3 Thin sections through some Triassic Tritrabidae showing test
structure and/or microsphere. aPaurinella sp. A, equatorial section,
scale bar =54 lm. bPaurinella aequispinosa Kozur & Mostler,
equatorial section, lower Ladinian, Rc4, scale bar =54 lm. cPau-
rinella sp. B, equatorial section, lower Ladinian, BV 85-70, scale
bar =42 lm. dAngulopaurinella edentata Dumitrica & Tekin n. sp.,
equatorial section, lower Ladinian, BV 85–70, scale bar =54 lm.
eZhamojdasphaera latispinosa Kozur & Mostler, transversal section
to microsphere, lateral view, 04-ELB-1, scale bar =54 lm. fSame,
detail of the central part of test showing one of the interradial pores of
the microsphere in the very centre, scale bar =11 lm. gSame
species and sample, equatorial section, scale bar =55 lm. hPara-
paronaella parvispinosa (Kozur & Mostler), equatorial section,
04-ELB-1, scale bar =42 lm. iRopanella zapfei (Kozur & Mostler),
equatorial section, 04-ELB-1, scale bar =42 lm. jTriassoastrum
noricum (Kozur & Mock), equatorial section, Glenn Shale, Alaska,
scale bar =58 lm. kSame, much magnified showing the triangular
microsphere, scale bar =11 lm. lParaparonaella robusta (Kozur &
Mostler), equatorial section, 04-ELB-1, scale bar =66 lm. mPseud-
angulobracchia prisca Dumitrica & Tekin n. sp., equatorial section,
04-ELB-1, scale bar =42 lm. nParaparonella okuyucui Dumitrica
& Tekin n. sp., transversal section through an arm, 04- ELB-1, scale
bar =21 lm. oTriassoastrum noricum (Kozur & Mock), transversal
section through an arm, Glenn Shale, Alaska, scale bar =21 lm.
pPseudangulobracchia prisca Dumitrica & Tekin n. sp., transversal
section through an arm, 04-ELB-1, scale bar =21 lm
b
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 355
123
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Fig. 4 a Paurinella curvata Kozur & Mostler, Rc4, scale bar =100
lm. bPaurinella mutihaci Dumitrica & Tekin n. sp., holotype, R78-
6, scale bar =115 lm. c–fPaurinella triangularis Kozur & Mostler:
cBR910, d–fR78-6, scale bar for all specimens =120 lm.
gPaurinella triassica (Kozur & Mostler), 04-ELB-2, scale
bar =90 lm. h,iPaurinella trifusus Dumitrica & Tekin n. sp.
hholotype, 04-ELB-1; iparatype, 04-ELB-2, scale bar for both =45
lm. j,kNeopaurinella ladinica Kozur & Mostler, both from Rc4,
scale bar =170 lm. lAngulopaurinella arborea Kozur & Mostler,
R78-6, scale bar =90 lm. mAngulopaurinella carpatica Dumitrica
& Tekin n. sp., holotype, R78-6, scale bar =110lm. n,oAngulop-
aurinella dentispinosa Dumitrica & Tekin n. sp.: nholotype, 04-ELB-1,
oparatype; both from 04-ELB-1, scale bar =90 lm. p–rAngulop-
aurinella edentata Dumitrica & Tekin n. sp.: pholotype, Rc4,
q,rparatypes, all from Rc4, scale bar for all specimens =85 lm.
s–wZhamojdasphaera aspinosa Kozur & Mostler: s-u04-ELB-1,
vR78-6, wBR908; scale bar for all specimens =70 lm
356 P. Dumitrica et al.
123
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Range and occurrence: Middle Longobardian, Spon-
goserrula rarauana subzone of the Muelleritortis cochle-
ata Zone, sample R78-6, Rarau Mt., Eastern Carpathians,
Romania.
Paurinella triangularis Kozur & Mostler 2006
(Fig. 4c–f)
2006 Paurinella triangularis Kozur & Mostler: 33, pl. 6
Figs. 1,2,5
Remarks: No internal structure could be found in this
species because of the poor preservation. Regardless, it
should be similar to that of the other species of the genus
Paurinella.
Range and occurrence: Late Ladinian (Longobardian)
from the Dinarides, Bosnia-Herzegovina; Eastern Carpa-
thians, Romania; and Zulla Formation, Wadi Bani Khalid,
Oman.
Paurinella triassica (Kozur & Mostler 1979)
(Fig. 4g)
1979 Spongotripus triassicus Kozur & Mostler: 79, pl.
11 Fig. 7, pl. 14 Fig. 9
Remarks: We have a single specimen from this species.
Its spines resemble the holotype and paratype of the spe-
cies, but the sides of its triangular test are less convex.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling, Austria; very rare in the
Ko
¨seyahya fauna, eastern Turkey.
Paurinella trifusus Dumitrica & Tekin n. sp.
(Fig. 4h, i)
Description: Shell triangular in face view, inflate, with
straight or slightly convex sides and three spines at corners.
Spines circular in transverse section, thick, and spindle-
shaped with slightly blunt tips, and about as long as the ray
of shell. Spongy shell with dense, polygonally framed
pores.
Material: Three specimens in samples 04-ELB-1 and
04-ELR-2, of which only two were photographed.
Holotype: Fig. 4h, Ko
¨seyahya section, Muse
´edeGe
´ol-
ogie Lausanne, No. MGL 96815.
Dimensions (in lm, based on two specimens): Length of
sides of spongy skeleton 94–100, length of spines 48–50.
Remarks: This species differs from the early late
Longobardian species Paurinella triangularis Kozur &
Mostler by having shorter and slightly blunt spines.
Etymology: From the Latin tri—three and fusus—spin-
dle, due to the spindle-shaped spines.
Range and occurrence: Very rare in the Middle Car-
nian, Tetraporobrachia haeckeli Zone, Ko
¨seyahya section,
eastern Turkey.
Paurinella spp. (A, B)
(Fig. 3a, c)
Remarks: These species are only illustrated to prove the
diversity of the structure of the spongy shell in this genus.
Occurrence: Latest Anisian or early Ladinian, Southern
Alps, North Italy.
Genus Neopaurinella Kozur & Mostler 1981
Type species:Neopaurinella sevatica Kozur & Mostler
1981.
Remarks: Given the asymmetry of the spongy shell and
the presence of a small spine in the vicinity of one normal
spine, it is possible that this genus came from the genus
Tetrapaurinella Kozur & Mostler by the reduction of one
spine. If this is the situation, Neopaurinella could be con-
sidered as ‘‘aborted Tetrapaurinella’’, and should be
assigned to the subfamily Tetrapaurinellinae n. subfam.
Unfortunately, we could not see the microsphere of the
single specimen we sectioned because of the poor preser-
vation of the central part of test.
Range: Early Ladinian or latest Anisian to early Norian.
Neopaurinella ladinica Kozur & Mostler 1994
(Fig. 4j, k)
1994 Neopaurinella ladinica Kozur & Mostler: 75, pl. 15
Figs. 5,6; pl. 16 Fig. 6
Remarks: Microsphere unknown because of its poor
preservation in the specimen sectioned.
Range and occurrence: Early Ladinian, Southern Alps,
North Italy.
Genus Spinopaurinella Kozur & Mostler 2006
Type species:Spinopaurinella magnispinosa Kozur &
Mostler 2006
Remarks: We found no species of this genus in our
material to investigate the microspheric structure.
Regardless, the spongy shell, the presence of three coplanar
spines, and the symmetry of shell suggest that the micro-
sphere is of tritrabid type.
Range: Longobardian of West Tethys.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 357
123
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Genus Angulopaurinella Kozur & Mostler 2006
Type species:Angulopaurinella crassa Kozur & Mostler
2006
Diagnosis: Globular or lenticular spongy spumellarian
radiolarians with three 3-bladed spines originated in the
corners of a tritrabid microsphere. Inside the test, spines
have several pairs of two opposite branches in successively
perpendicular planes, on the proximal portion, and irregu-
larly disposed branches on middle and distal portions.
Remarks: The microsphere of this genus, studied in both
the type species and in some undescribed late Illyrian or
Fassanian species (Fig. 1b, g), is of tritrabid type and
similar to that of Paurinella Kozur & Mostler and
Zhamojdasphaera Kozur & Mostler, having three corners
in the equatorial or subequatorial plane, three lateral pores,
one on each side between corners, and a three-armed
structure on each face. This structure is usually a little
complicated by the appearance of additional bars, and
disturbed by the bars arising from it to build the perimi-
crospheric skeleton. Unlike the late Ladinian and Carnian
species, the late Illyrian and Fassanian species have non-
serrate and usually untwisted or slightly twisted spines.
Range: Late Illyrian or early Ladinian to middle Carnian
so far as known.
Angulopaurinella arborea Kozur & Mostler 2006
(Fig. 4l)
2006 Angulopaurinella arborea Kozur & Mostler: 31, pl.
4 Fig. 2, pl. 9 Fig. 5
Remarks: The illustrated specimen is the oldest speci-
men of this genus so far described. Based on the mor-
phology of its spines, where the three-bladed character
is very slightly marked, we consider, in agreement with
Kozur and Mostler (2006), that this species is one of the
transitional taxa between Spinopaurinella Kozur & Mostler
and Angulopaurinella.
Range and occurrence: Middle to early late Longobar-
dian, Muelleritortis cochleata Zone, Spongoserrula
rarauana and S. fluegeli Subzones: Dinarides, Bosnia-
Herzegovina; Rarau Mt., eastern Carpathians, Romania.
Angulopaurinella carpatica Dumitrica & Tekin n. sp.
(Fig. 4m)
Description: Test flat, subcircular with superficial nodes
interconnected by thin bars forming triangular meshes.
Spines three-bladed, slightly longer than test ray, and about
360°dextrally twisted. They increase slightly in thickness
distally, on the proximal half, then decrease fast and ter-
minate in a pointed vertex. Blades of spines thick with
rounded outer margin. Interradial periphery of test may
have one, or likely more, short and thin secondary spines.
Material: A single specimen (the holotype) in sample
R78-6.
Holotype: Fig. 4m, Muse
´edeGe
´ologie Lausanne, No.
MGL 96816.
Dimensions (in lm based on holotype): Diameter of test
164, length of spines 117, maximum diameter of spines 48.
Remarks: Although based on a single specimen, this
species is very well distinguished from all the other species
of this genus in having well-twisted spines, flat shell, and
nodose surface.
Etymology: From its occurrence in the Carpathian
Mountains.
Range and occurrence: Late Ladinian (middle Longo-
bardian), Muelleritortis cochleata Zone, Spongoserrula
rarauana Subzone, sample R78-6, base of Piatra Zimbrului
olistolith, Rarau Mt., Eastern Carpathians, Romania.
Angulopaurinaella dentispinosa Dumitrica & Tekin n. sp.
(Fig. 4n–o)
Description: Spongy test lenticular, triangular in face
view. Spongy meshwork massive or slightly organized in
about three to four weakly visible layers. Spines robust,
three-bladed, sinistrally twisted up to 90°-180°. They
increase slightly in thickness up to the middle part, and
slowly decrease distally. Blades bear one to three low,
broad teeth or very short thorns.
Material: Twenty specimens in sample 04-ELB-1, of
which three have been photographed.
Holotype: Fig. 4n, Muse
´edeGe
´ologie Lausanne, No.
MGL 96817.
Dimensions (in lm, based on 3 photographed speci-
mens): Diameter of shell 77–105, length of spines 85–89.
Remarks: The species very much resembles the Longo-
bardian species A. crassa Kozur & Mostler and A. spiralis
Kozur & Mostler, from which it differs by having very
small teeth on spines and blades strongly sinistrally twis-
ted. The spines of A. spiralis are also twisted, but its
holotype shows a variable twist, one spine is sinistrally
twisted, another one is dextrally twisted, and the third
seems to have also a slight dextral twisting.
Etymology: From the Latin dens, -tis—tooth and spino-
sus, -a, -um—spiny.
358 P. Dumitrica et al.
123
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Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, east of Elbistan,
eastern Turkey.
Angulopaurinella edentata Dumitrica & Tekin n. sp.
(Figs. 1h, 3d,4p–r)
Description: Test small, inflate lenticular, spongy,
irregularly organized, and slightly triangular in face view,
with three coplanar three-bladed spines. Spines pointed,
gently narrowing distally, with blades simple, straight or more
or less dextrally twisted, especially on the distal part. Test
multi-layered with small polygonal meshes of various sizeand
irregular arrangement on the surface. Microsphere as with the
genus, with asymmetric faces, one simple with a three-barred
structure of Tritbabs-type, the other one more complex, with a
small three-barred structure in the face centre, three pores
aligned on the direction of the three primary beams and three
pores on the direction of the three lateral equatorial pores.
Material: Ten specimens in sample Rc4, of which three
were photographed.
Holotype: Fig. 4p, Muse
´edeGe
´ologie Lausanne, No.
MGL 96818.
Dimensions (in lm, based on three photographed speci-
mens): Diameter of spongy test 71–119, length of spines
94–119.
Remarks: This new species differs from the Longobar-
dian and Carnian species of the genus Angulopaurinella so
far described in having no denticles on the blades of
the spines and in being less twisted. The variability of the
species is mainly expressed in the twisting degree of the
spines. The specimens with untwisted spines are less fre-
quent than those with twisted spines.
Etymology: From the Latin edentatus, -a, -um—without
teeth.
Range and occurrences: Latest Ansian (late Illyrian) or
early Ladinian from Recoaro (Vicentinian Alps, North
Italy) and Marmolada Massif (Dolomites, North Italy).
Genus Zhamojdasphaera Kozur & Mostler 1979
Type species:Zhamojdasphaera latispinosa Kozur &
Mostler 1979.
Diagnosis: Test spherical to subspherical, spongy, many-
layered with three broad, foliaceous spines originated in a
tritrabid microsphere. Spines straight or twisted, and flat-
tened in vertical planes.
Remarks: The genus Zhamojdasphaera was described
only on the basis of the type species, and was initially
included within the Stylosphaerinae and later questionably
within the Oertlispongidae. Later, Kozur and Mostler
(2006) described several new Longobardian species and
erected a special family for it. As mentioned above under
the subfamily Intermediellinae, it is not necessary to erect
a special family for this genus only because its spines are
foliaceous. The peculiarity of its spines appeared first in
the late Ladinian (Longobardian), practically simulta-
neously with the advanced foliaceaous Oertlispongidae,
and we consider it to be the result of a lateral gene transfer
from the foliaceous Oertlispongidae to a three-spined
Ladinian spongy radiolarian (Dumitrica and Guex
2003). Such a radiolarian should have been a species of
Paurinella Kozur & Mostler. In agreement with Kozur
and Mostler (2006), the most primitive species of
Zhamojdasphaera is Zh. transita Kozur & Mostler, from
the lower part of the upper Longobardian. This species
differs from the other species of Paurinella in having only
two very narrow wing-like lateral thorns within the inner
third of the primary spines.
Until present, no information was given on the type of
microsphere, although Moix et al. (2007,pl.4Figs.6a,
6b) showed a remnant of it, but where no special struc-
ture could be seen, because the specimen was illustrated
in lateral view and also because the optic microscope is
usually better than the electronic microscope for deci-
phering the structure of the microsphere of the spongy
radiolarians. In the centre of the spongy shell, this
specimen shows only an incomplete frame of one lateral
or interradial pore of the microsphere (see also Fig. 3e, f
in the present article). The sections we made in the very
well preserved specimens of the type species have pro-
ven that the skeleton has five to nine layers of closely
spaced shells and that in its centre is a microsphere of
tritrabid type (Figs. 1f, 3g). Like any microsphere of this
type (De Wever et al. 2001, Fig. 85.4–5), it is triangular
in axial view, has three flattened corners in the equatorial
plane, one interradial pore in the middle of each side, and
a three-armed structure in the middle of one or both
faces.
The genus began with straight, flattened spines
approximately in the middle Longobardian, as show the
species illustrated by Kozur & Mostler (2006), and at
approximately the same level, the spines started twisting
dextrally or sinistrally. The sense of twisting is probably a
specific feature, as it is for the species of other genera. Species
with untwisted spines continued, however, to co-exist with
those with twisted spines until the early Norian.
Range: Late Ladinian (Longobardian) to early Norian
(Lacian).
Zhamojdasphaera aspinosa Kozur & Mostler 2006
(Fig. 4s–w)
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 359
123
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2006 Zhamojdasphaera aspinosa Kozur & Mostler: 36,
pl. 8 Fig. 3
Remarks: The Longobardian specimens we assign to this
species are perfectly similar to the holotype, whereas some
middle Carnian specimens have shorter and wider spines.
Although these specimens may be assigned to a new species, in
the present article we include them in Z. aspinosa, because of
their untwisted spines and the absence of distal needle-like thorns.
Range and occurrence: Late Ladinian to middle Car-
nian; late Ladinian (middle Longobardian), Rarau Mt.,
eastern Carpathians, Romania; early late Longobardian of
Dinarides, Bosnia-Herzegovina; middle Carnian, Ko
¨seya-
hya section, Elbistan, SE Turkey.
Zhamojdasphaera blechschmidti Dumitrica & Tekin n. sp.
(Fig. 5a, b)
Description: Test spongy, globular as for the genus.
Spines flat in vertical planes, foliaceous, non-twisted, ogee-
shaped in lateral view, and terminated in a short spine.
Both lateral margins of spines very broad forming two
longitudinal ridges separated by a longitudinal groove.
Material: One specimen in BR913.
Holotype:Fig.5a, b, sample BR913, Wadi Bani Khalid
section, Oman, Muse
´edeGe
´ologie Lausanne, No. MGL 96819.
Dimensions (in lm based on holotype): Diameter of
shell 100, length of spines 80, breadth of spines 70.
Remarks: Although we have a single incomplete speci-
men of this species, the morphology of its spines differs
from that of all the other species of this genus. A single
known species with spines having thickened margins is Zh.
furcata n. sp., but this species has forked spines, and its
margins are not divided into two ridges as Zh. ble-
chschmidti n. sp.
Etymology: The species is dedicated to Dr. Ingo Ble-
chschmidt, NAGRA, Switzerland, for his contribution to
the geology of the Hawasina Complex, Oman.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Zulla Formation, Wadi Bani Khalid
section, Oman.
Zhamojdasphaera furcata Dumitrica & Tekin n. sp.
(Fig. 5c, d)
Description: Shell spongy, globular. Spines short,
untwisted, broad in vertical plane, ogee-shaped, and ter-
minated into two short spines. Margins of spines thickened.
Material: One specimen in BR913, Wadi Bani Khalid
section, Oman.
Holotype: Fig. 5c, d, Muse
´edeGe
´ologie Lausanne, No.
MGL 96820.
Dimensions (in lm based on holotype): Diameter of
shell 105, length of spines 97, breadth of spines 75.
Remarks: This new species differs from the other spe-
cies of the genus in having forked spines.
Etymology: From the Latin furcatus, -a, -um—forked.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Zulla Formation, Wadi Bani Khalid
section, sample BR913, Oman.
Zhamoidasphaera goricanae Dumitrica & Tekin n. sp.
(Fig. 5e, f)
1990 Zhamojdasphaera sp. Gorican & Buser: 161, pl. 2
Fig. 9
Description: Test triangular in face view with a rela-
tively thick cortical shell with small and irregularly
arranged pores. At corners, this test passes gradually into
the foliaceous spines. Inner shell unknown. Spines flat in
vertical planes proximally, thin, and twisted dextrally in the
middle part up to 70–90°and broadened. Distal part lan-
ceolate with straight or slightly concave sides.
Material: Four specimens in sample R78-6, of which
two have been photographed.
Holotype: Fig. 5e, Muse
´edeGe
´ologie Lausanne, No.
MGL 96821.
Dimensions (in lm based on four specimens): Length of
triangular sides of shell 81–89, of spines 113–121.
Remarks: This species differs from all the other species
of the genus especially by having a triangular shell and a
gradual transition between shell and spines. The specimens
from the type sample (R78-6) differ from the specimen
illustrated by Gorican and Buser (1990) from the Longo-
bardian of Slovenia, in having broader spines, but have in
common with this specimen the triangular morphology of
the shell.
Etymology: The species is named for Dr. Spela Gorican,
Slovenia, who illustrated a specimen of this species for the
first time.
Range and occurrence: Lower to middle Longobardian:
Vrsic, Slovenia, and Rarau Mt., Eastern Carpathians, Romania.
Zhamojdasphaera komoi Dumitrica & Tekin, n. sp.
Fig. 5g–j
1979 Zhamojdasphaera latispinosa Kozur & Mostler,
part: 67, pl. 7 Fig. 7,8,9, non pl. 12 Fig. 5
360 P. Dumitrica et al.
123
Author's personal copy
Fig. 5 a–bZhamojdasphaera blechschmidti Dumitrica & Tekin, n.
sp., holotype, BR913, scale bar =130 lm. c,dZhamojdasphaera
furcata Dumitrica & Tekin, n. sp. holotype, BR913, scale bar =90
lm. e,fZhamojdasphaera goricanae Dumitrica & Tekin n. sp.
eholotype, fparatype, both from R78-6, scale bar =120lm. g–j
Zhamojdasphaera komoi Dumitrica & Tekin n. sp. gholotype,
04-ELB-1, scale bar =120 lm; h-jparatypes, all from 04-ELB-2,
scale bar =120 lm. k,mZhamojdasphaera latispinosa Kozur &
Mostler: kand l04-ELB-1, m04-ELB-2, scale bar for all
specimens =120 lm. nZhamojdasphaera cf. rigoi Kozur & Mostler,
04-ELB-2, scale bar =120 lm. o,pZhamojdasphaera robinsoni
Dumitrica & Tekin, n. sp.: oholotype, pparatype, both from Glenn
Shale, scale bar =150 lm. qTritrabs worzeli (Pessagno), early
Tithonian, Solnhofen area, Germany, scale bar =250 lm. r–t
Triassoastrum baumgartneri Dumitrica & Tekin n. sp.: rholotype,
04-ELB-2, s,tparatypes, both from 04-ELB-5; scale bar for all
specimens =120 lm. uTriassoastrum noricum (Kozur & Mock),
Glenn Shale, scale bar =200 lm. v,wTriassoastrum paratrammeri
Dumitrica & Tekin n. sp.: vholotype, wparatype, both from
04-ELB-2, scale bar =230 lm
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 361
123
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Description: Test globular, slightly flattened axially with
subtriangular outline. Pores small, polygonally framed and
irregularly arranged. Spines slightly longer than the
diameter of shell and well flattened. Their proximal half
straight and their distal half usually up to 90°dextrally
twisted, narrowing distally, and pointed.
Material: Four photographed specimens in the Ko
¨seya-
hya section (samples 04-ELB-1 and 04-ELB-2).
Holotype: Fig. 5g, Muse
´edeGe
´ologie Lausanne, No.
MGL 96822.
Dimensions (in lm based on 4 specimens): Diameter of
shell 83–89, length of spines 112–126.
Remarks:Zhamojdasphaera komoi n. sp. differs from
Zh. latispinosa Kozur & Mostler in having the proximal,
untwisted portion of spines longer and the distal portion
less twisted.
Etymology: The species is named for Dr. Heinz Kozur
and Prof. Dr. Helfried Mostler who, in their paper pub-
lished in 1979, illustrated three specimens of this species as
paratypes of Zhamojdasphaera latispinosa.
Range and occurrence: Middle Carnian, Tetraporobrachia
haeckeli Zone: Go
¨stling, Austria; Ko
¨seyahya section, eastern
Turkey; latest Tuvalian?- early Lacian, Glenn Shale, Alaska.
Zhamojdasphaera latispinosa Kozur & Mostler 1979
(Fig. 5k, m)
1979 Zhamojdasphaera latispinosa Kozur & Mostler:
67, pl. 12 Fig. 5, non pl. 7 Figs. 7–9;
1984 Zhamojdasphaera latispinosa Kozur & Mostler.—
Lahm: 74, pl. 13 Fig. 5
1990 Zhamojdasphaera latispinosa Kozur & Mostler.—
Gorican & Buser: 161, pl. 2 Fig. 8
1999 Zhamojdasphaera latispinosa Kozur & Mostler.—
Tekin: 68, pl. 2 Fig. 13
Material: Several tens of specimens.
Remarks: The sections in the shell show that the shell
consists of four to six equidistant perimicrospheric layers
interconnected by numerous radial bars, which usually do
not continue from one interlayer to another. The spines of
this species in our samples and in the literature are dex-
trally twisted, and they show a certain variability of the
length of the proximal untwisted portion.
Range and occurrence: Late Ladinian (Longobardian)?
to Middle Carnian (Julian): Longobardian? from central
Slovenia; middle Carnian from Go
¨stling and Grossreifling,
Austria; Bozkir, Konya, and Ko
¨seyahya section, Elbistan,
SE Turkey; chert member of Zulla Formation, Wadi Bani
Khalid, Oman.
Zhamojdasphaera cf. rigoi Kozur, Moix & Mostler in Moix
et al. 2007
(Fig. 5n)
cf. 2007 Zhamojdasphaera rigoi Kozur, Moix & Mostler
in Moix et al.: 297, pl. 4 Figs. 2, 4–6
Remarks: Our specimens, very rare, differ from the
holotype of the species by having spines with a much more
rounded margin in face view. By this character, they have a
transitional position between Zh. rigoi and Zh. latispinosa.
In fact, the differences between the two species are very
small. The differences between the two subspecies of Zh.
rigoi (Zh. rigoi rigoi and Zh. rigoi brevispinosa), which
Kozur et al. (2007) discriminated in the sample G11 from
the Mersin Me
´lange, are also very small. The latter sub-
species should be considered a morphological variety of
Zh. rigoi without a formal status.
Range and occurrence: Middle Carnian, upper part of the
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section, Elbis-
tan, SE Turkey; early late Carnian (early Tuvalian), Spongo-
tortilispinus moixi Zone, Mersin Me
´lange, SE Turkey.
Zhamojdasphaera robinsoni Dumitrica & Tekin n. sp.
(Fig. 5o, p)
Description: Test rounded triangular in face view, with
flattened faces. Cortical layer of test of pseudoaulophacid
structure, consisting of nodes interconnected by bars forming
triangular meshes. Spines as long as diameter of shell and
twisted dextrally 90°. They terminate in a short axial thorn and
bear two pointed lateral corners resulting from a deep undu-
lation of the blades. Between these corners and the distal thorn
the margin of spines is concave in polar projection.
Material: Seven specimens in the Glenn Shale, east-
central Alaska, three of which have been photographed.
Holotype: Fig. 5o, Muse
´edeGe
´ologie Lausanne, No.
MGL 96823.
Dimensions (in lm based on three specimens) Diameter
of shell 115–135, length of spines 126–128.
Remarks: This new species differs from the other species
of the genus in having a shell with pseudoaulophacid cortical
structure and flat faces, and in the morphology of spines.
Etymology; The species is dedicated to Dr. B.F. Robin-
son for his unpublished contribution to the upper Triassic
radiolarian biostratigraphy of the Glenn Shale, Alaska.
362 P. Dumitrica et al.
123
Author's personal copy
Range and occurrence: Uppermost Tuvalian (upper-
most Carnian) or lower Lacian (lowermost Norian), Glenn
Shale, East-Central Alaska.
1b. Subfamily TITRABINAE Baumgartner 1980
Type genus:Tritrabs Baumgartner 1980
Remarks: At present, when we know the microsphere
and the arm structure of the Triassic three-armed taxa
assigned to the Tritrabidae, we can see that the members of
this group represent a rather unitary and monophyletic
taxonomic unit. Although no transition has been recorded
so far, the stratigraphic range and the microspheric struc-
ture suggest that the subfamily has its origin in the genus
Paurinella by the extension of the spongy test along the
three primary beams, forming spongy arms, and reorgani-
zation of this test to create an internal symmetry. During
this process, the microsphere simplified and increased its
symmetry, always preserving the fundamental elements
described and illustrated in De Wever et al. (2001).
Range: Late Triassic (Carnian) to Late Cretaceous
(Maastrichtian).
Genus Triassoastrum Kozur & Mostler 1978, emended
herein
Type species:Triassoastrum transitum Kozur & Mostler
1978 =Rhopalodictyum glaber Kozur & Mostler 1978.
1978 Triassoastrum n. gen. Kozur & Mostler: 154
1981 Paratriassoastrum n. gen. Kozur & Mostler: 63
Emended diagnosis: Three-armed test with a micro-
sphere of tritrabid type and arms with a primary beam, six
secondary beams inside arms, and up to 12 tertiary or
cortical beams that can be continuous or interrupted, or
absent, in which case they are replaced by a rather irregular
system of polygonal frames. When cortical beams are
continuous, cortical shell has pores in square patters
aligned in 12 longitudinal rows separated by the 12 beams
and by numerous circumferential bars. On each level,
transversal sections of arms have 12 pores, of which three
are primary pores, three are secondary pores and six are
tertiary pores (Fig. 2c). Pores on each layer alternate with
pores of neighbouring layer so that there are no visible
canals along arms. Central part of test very small and flat,
without patagium.
Remarks: The genus Triassoastrum Kozur & Mostler
was described for a poorly illustrated specimen coming
from the middle Carnian Reiflinger Limestone from Go
¨s-
tling, Austria. According to the original diagnosis and
illustration of the type species (T. transitum Kozur &
Mostler), the only species described in this genus so far and
very probably the only specimen recorded so far, this genus
differs from the other three-armed species of Carnian
radiolarians in having one arm distally forked in a plane
oblique to the equatorial plane of the test. The diagnosis
specifies also that the arms have pores arranged irregularly
or in longitudinal rows. The latter character is rather dif-
ficult to see on the figure of the holotype (Kozur and
Mostler 1978: 154, pl. 3 Figs. 6–7), but what can be easily
seen on one of its arms is the arrangement of pores in
transversal rows. This proves that the structure of its
arms is similar to that of T. noricum (Kozur & Krahl),
T. trammeri (Kozur & Mostler) and other Late Triassic
three-armed species. The morphological character Kozur
and Mostler (1978) used to distinguish this genus (one
forked arm) is insignificant. In fact, the holotype of the type
species is clearly an anomaly of Rhopalodictyum glaber
Kozur and Mostler 1978, pl. 3 Fig. 15) with a small arm
grown on one of the normal arm of the specimen. This
phenomenon is not uncommon among armed radiolarians.
For comparison, we illustrate herein similar anomalies of
an early Tithonian specimen of Tritrabs worzeli (Pessagno)
from the Solnhofen area, Germany (Fig. 5q). In this situ-
ation, R. glaber becomes a synonym of Triassoastrum
transitum Kozur & Mostler and the genus, in order to keep
it, is necessary to be emended to comprise exclusively
three-armed Triassic radiolarians. It is what we have done
in this article.
The genus Paratriassoastrum Kozur & Mostler should
be considered a younger synonym of Triassoastrum Kozur
& Mostler because its type species—P. austriacum Kozur
& Mostler—has all characters of the type species of the
latter genus; namely, its additional arm does not arise from
the centre of the shell, but obliquely from one arm. The
only difference between the two type species of the two
genera is that in T. transitum the additional arm grows from
the outer third of the arm, whereas with P. austriacum it
grows from the inner third. However, the distance from the
centre from where an arm forks cannot represents a generic
character. Even for a specific character this distance is
difficult to use for three-armed or four-armed species with
forked arms. In fact, it is difficult to understand why Kozur
and Mostler (1981) chose P. austriacum as type species
when they erected the genus Paratriassoastrum, because in
the same paper they described a P. cordevolicum Kozur &
Mostler that has all four arms originated in the centre. In
addition to T. transitum, this species (P. austriacum)
should be also considered an anomaly of a species with
three arms. Such a species seems to be Rhopalodictyum
simoni Kozur & Mostler that very much resembles P.
austriacum and that was described from the same middle
Carnian fauna.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 363
123
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Externally, the cortical structure of the arms of the type
species of Triassoastrum resembles that of the type species
of Homoeoparonaella Baumgartner in having the cortical
shell with square pore frames arranged in longitudinal and
transversal rows, usually with nodes at corners. However,
whereas in the latter genus the nodes are equal [see H.
elegans (Pessagno) in Baumgartner 1980], they are of two
sizes in this new genus, along each external beam existing a
regular repetition of smaller and bigger nodes. Laterally, the
bigger nodes along one longitudinal row correspond to
smaller nodes of the neighbouring longitudinal rows and
vice versa (Fig. 6j). In the literature, this feature is very
clearly visible in Triassoastrum noricum (Kozur & Mock)
illustrated by Tekin (1999)asParonaella norica.This
difference in cortical structure between the two genera
represents an external reflection of the internal arm struc-
ture. According to the illustration by Baumgartner (1980,
pl. 1 Fig. 15, pl. 2 Figs. 5, 6, 15, 16) and Nagai (1985), the
arms of Homoeoparonaella Baumgartner have three pri-
mary canals around the primary beam, surrounded by six or
nine secondary canals. This means that along the primary
beam, the three branches arise always in the same plane. In
Triassoastrum, the situation is different (Fig. 2b); due to the
permanent inversion of the branches of the primary and
secondary beams there is no canals, because in axial pro-
jection of arms the pores of a layer are closed by the triradial
connections among the beams of the neighbouring layers.
Externally, not all species included herein in this genus
have pores arranged in a square pattern. In some species,
this structure is covered by a cortical layer with disorderly
arranged pores (see Triassoastrum paratrammeri n. sp.,
Figs. 5v, w, 6a, b).
Microsphere of Triassoastrum resembles that of Tritrabs
and other Jurassic and Cretaceous tritrabids in its general
structure, having three corners in the equatorial plane, one
interradial pore on each side between corners, and a three-
armed structure on each face (Figs. 1d, e, 2a). This struc-
ture is usually disturbed by the appearance of additional
bars among the frames of the interradial pores, a character
that was never found in the Jurassic and Cretaceous tri-
trabids that show a simplification and stabilization of the
microsphere. On the proximal portion of the primary beams
the first branches have an irregular disposition, then they
start gradually to be disposed in verticils of three branches
lying in successively inverted planes (Fig. 2a), the result
being the absence of canals along arms as said above.
Sometimes the verticils are disturbed, or there are no
verticils, the three branches originating at different levels
along the primary beam.
Range: Middle Carnian to early Norian or even Rhaetian
if the specimen illustrated by Carter (1993, pl. 10 Fig. 9)as
Paronaella sp. cf. pacofiensis Carter belongs to this genus.
Triassoastrum baumgartneri Dumitrica & Tekin n. sp.
(Fig. 5r–t)
Description: Test structure as of the genus. Arms long,
circular to subcircular in cross-section with about 12 cor-
tical longitudinal beams intersected by 10–12 circumfer-
ential bars to form rows of pores in square pattern. Tips of
arms expanded with one long axial spine and two thin and
shorter lateral spines. Base of axial spine conical. Pores of
tips smaller than of arms and usually disorderly arranged.
Material: Seven specimens in samples 04-ELB-2 and
04-ELB-5, of which three photographed.
Holotype: Fig. 5r, Muse
´edeGe
´ologie Lausanne, No.
MGL 96824.
Dimensions (in lm, based on three specimens): Length
of arms without spines 228–260, width of arms at base
50–60, length of distal spines 70–140.
Remarks: This species resembles T. trammeri (Kozur &
Mostler) in cortical test structure from which it differs in
having expanded tips with one long axial spine and 2 thin
and smaller lateral spines. From Paraparonella okuyucui n.
sp. this species differs in having only three spines on tips,
central part of tips conical, cortical pores arranged in
square pattern, and well marked cortical longitudinal
beams.
Etymology: The species is named for Dr. P. O. Baum-
gartner, University of Lausanne, Switzerland, to honour his
great contribution to the taxonomy and biostratigraphy of
Mesozoic radiolarians.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Triassoastrum noricum (Kozur & Mock 1981 in Kozur &
Mostler 1981)
(Fig. 5u)
1981 Paronaella norica n. sp. Kozur & Mock.—Kozur
& Mostler: 61, pl. 46 Fig. 2
1999 Paronaella norica Kozur & Mock.—Tekin: 89, pl.
9 Figs. 9, 11
1999 Paronaella norica Kozur & Mock.—Bragin &
Krylov: 554, Figs. 8g–h
2007 Paronaella norica Kozur & Mock.—Bragin: 1001,
pl. 12 Fig. 1
Remarks: According to the original description, this
species resembles T. trammeri (Kozur & Mostler) from
which it differs in having club-shaped tips. By its very well
visible cortical structure with pores in square pattern, it is
364 P. Dumitrica et al.
123
Author's personal copy
Fig. 6 a,bTriassoastrum paratrammeri Dumitrica & Tekin n. sp.,
paratypes: a04-ELB-5, bBR918, scale bar for both speci-
mens =230 lm. cTriassoastrum cf. trammeri (Kozur & Mostler),
04-ELB-2, scale bar =80 lm.d,eTriassoastrum simplicissimum
Dumitrica & Tekin n. sp.: dholotype, 04-ELB-5; eparatype, 04-ELB-
3; scale bar for both specimens =90 lm. f–lTriassoastrum trammeri
(Kozur & Mostler): f–h04-ELB-2, i04-ELB-1, j04-ELB-5, kGlenn
Shale, ldetail of the cross section in an arm 04-ELB-3, scale
bar =160, 160, 160, 160, 70, 160 and 70 lm, respectively.
m,nParaparonaella claviformis (Kozur & Mostler): m04-ELB-1,
N04-ELB-4, scale bar for both specimens =250 lm. o,pParapa-
ronaella fragilis (Kozur & Mostler), 04-ELB-1, scale bar =230 lm.
q–uParaparonaella okuyucui Dumitrica & Tekin n. sp.: qholotype,
04-ELB-2, scale bar =190 lm; r–uparatypes: q04-ELB-1, r–u04-
ELB-2, udetail of arm structure of Fig. t;scale bar =190, 190, 190
and 60 lm, respectively. v–wParaparonaella parvispinosa (Kozur &
Mostler): v04-ELB-2, w04-ELB-5, scale bar =230 lm. xParapa-
ronaella robusta (Kozur & Mostler), 04-ELB-5, scale bar =240 lm
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 365
123
Author's personal copy
the closest taxon to this species, from which it might have
occurred by the disappearance of the terminal spine and
development of a cap-like spongy structure. As mentioned
above under the genus, the peculiarity of the arm structure
of this genus is best emphasized in the specimens illus-
trated by Tekin (1999, pl. 9 Fig. 9and especially Fig. 11).
Range and occurrence: Latest Carnian (late Tuvalian)
to early Norian (Lacian): Sulov village, West Carpathians,
Slovakia; Antalia, Turkey; southern Cyprus; Zula Forma-
tion, Wadi Bani Khalid, Oman; Glenn Shale, Alaska.
Triassoastrum paratrammeri Dumitrica & Tekin n. sp.
(Figs. 5v, w, 6a, b)
1978 Rhopalodictyum trammeri Kozur & Mostler (part):
152, pl. 3 Fig. 11
1984 Paronaella simoni Kozur & Mostler (part).—
Lahm: 89, pl. 16 Fig. 6, non 5
Description: Arms long, circular in cross-section,
increasing slightly in diameter distally, and terminating in a
cone the vertex of which is prolonged in a long needle-like
spine aligned with the primary beams. Surface of arms with
disorderly arranged polygonal to oval pore frames and
partly with square pore frames in longitudinal and trans-
versal rows, and all longitudinal beams of equal height.
Material: Eight specimens, five of them photographed,
in samples 04-ELB-2, 04-ELB-5 and BR918.
Holotype: Fig. 5v, Muse
´edeGe
´ologie Lausanne, No.
MGL 96825.
Dimensions (in lm, based on 5 photographed speci-
mens) Length of arms without spines 200–250, diameter
of arms at base 50–80.
Remarks: By its shape, this species very much resembles
T. trammeri (Kozur & Mostler), from which it only differs
in having all or only a part of cortical pore frames of arms
polygonal and disorderly arranged, by having arms slightly
spindle-shaped, perfectly circular in cross-section, and
usually longer distal spines.
Etymology: From the Latin para—besides to, closely
related to, and trammeri—a species name.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Grossreifling, Austria; Ko
¨seyahya
section, SE Turkey; Late Carnian (early Tuvalian),
Spongotortilispinus moixi Zone, Zulla Formation, Wadi
Bani Khalid section, Oman.
Triassoastrum simplicissimum Dumitrica & Tekin n. sp.
(Fig. 6d, e)
Description: Arms cylindrical with blunt tips and short
and thin axial distal spines. Surface of arms with five to six
visible longitudinal beams on half diameter which, on some
portions, some are higher than their neighbouring beams.
Transversal partitions well visible, in number of maximum
10–11 on each arm giving rise to pores arranged in a square
pattern. On mature specimens (Fig. 6d), this structure is
partly screened by a thinner superficial layer with oval or
subcircular pores of rather irregular arrangement.
Material: Three specimens in the Elbistan material.
Holotype: Fig. 6d, Muse
´edeGe
´ologie Lausanne, No.
MGL 96825.
Dimensions (in lm, based on three specimens) Length of
arms without distal spine 102–154, diameter of arms
38–46.
Remarks:T. simplicissimum n. sp. is morphologically
close to T. trammeri from which it differs in having a
smaller number of superficial longitudinal beams, a less
regular surface, and blunt tips.
Etymology: From the Latin meaning the simplest.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Triassoastrum trammeri (Kozur & Mostler 1978)
(Figs. 1d, e, 2a–c, 6f–l)
1978 Rhopalodictyum trammeri Kozur & Mostler: 152,
pl. 1 Figs. 3, 8; pl. 3, Figs. 8–11, 16
1981 Paronaella trammeri (Kozur & Mostler).—Kozur
& Mostler: 61
1990 Homoeoparonella sp. A.—Yeh: 12, pl. 7 Fig. 2
1999 Paronaella trammeri (Kozur & Mostler).—Tekin:
90, pl. 10 Fig. 4
Description: Initial structure of tritrabid type as with the
genus. Arms long, practically equal and equidistant, cir-
cular to subcircular in cross-section. They increase very
slightly in diameter distally and terminate in a conical tip,
with a commonly long, thin distal spine. Arms with an
axial primary beam bearing about 15 verticils of three
branches in successively inverted position, six secondary
beams forming a central tube, and about 12 cortical beams
interconnected by circumferential bars forming about 12
longitudinal rows of about 15 square pores. Tips of arms
conical, simple or, probably in old specimens, with an
additional layer forming large irregular meshes.
Material: About one hundred specimens. At this strati-
graphic level, it is the most frequent species of the genus.
366 P. Dumitrica et al.
123
Author's personal copy
Remarks: Although the species is characterized by
having arms with conical distal ends armed with a thin
axial spine, a single specimen was found to have three
distal spines on an arm (Fig. 6f).
Range and occurrence: Middle Carnian (Tetraporobra-
chia haeckeli Zone) to early Norian: Go
¨stling, Austria;
Kemer area, Antalya, and Ko
¨seyahya section, Elbistan, SE
Turkey; Zulla Formation, Wadi Bani Khalid section,
Oman; early Norian, Glenn Shale, Alaska.
Triassoastrum cf. trammeri (Kozur & Mostler 1978)
Fig. 6c
Remarks: The illustrated specimen, the only one of this
type we found, has half the number of longitudinal beams
and transverse partitions by comparison with T. trammeri,
is much smaller than it, and appears to be a young onto-
genetic stage of this species. If so, its cortical shell would
represent the intermediary shell of the arms of mature
specimens. And if this interpretation were correct, it would
mean that the true cortical shell of T. trammeri was built in
a last ontogenetic stage and very fast, judging by the rarity
of this morphotype by comparison with the high frequency
of mature specimens.
Range and occurrence: Very rare in the Ko
¨seyahya
section, middle Carnian, (Tetraporobrachia haeckeli
Zone).
Genus Paraparonaella Dumitrica & Tekin n. gen.
Type species:Rhopalodictyum reiflingensis Kozur &
Mostler 1978.
Diagnosis: Three-armed, usually flat tritrabids with the
skeleton of Triassoastrum type covered by a cortical
spongy layer of various thickness. Arm tips usually
expanded and provided with an axial spine and two or more
lateral spines or spongy projections.
Remarks: The genus Paraparonaella n. gen. resembles
the genus Paronaella Pessagno in shape and spongy test,
but differs structurally from it in having a tritrabid micro-
sphere. As already shown (De Wever et al. 2001,
Fig. 81.1–5), the microsphere of Paronaella is of angu-
lobrachiid type, having an eight-shaped outline. Based
on what we know now about the initial structure of the
Paronaella-like Triassic radiolarians, we consider that the
genus Paronaella is missing in the Triassic, or at least in
the Carnian and Norian, its first occurrence should be either in
the Rhaetian or in the lower Jurassic. Paraparonaella has the
same microspheric and core arm structures as Triassoastrum
Kozur & Mostler. In fact, it could be considered a Triassoa-
strum with arms surrounded by a spongy cortical shell and
usually bearing lateral projections or spines on arm tips.
Besides the species occurring in the Ko
¨seyahya samples,
Rhopalodictyum suborbiformis Kozur & Mostler and R.
hirsutum Kozur & Mostler can also be assigned to this
genus.
Etymology: From the Latin para—beside, closely rela-
ted to, and paronaella—a genus name. Feminine gender.
Range: Middle Carnian, Tetraporobrachia haeckeli Zone
so far as known.
Paraparonaella claviformis (Kozur & Mostler 1978)
(Fig. 6m, n)
1978 Rhopalodictyum claviformis Kozur & Mostler: 147,
pl. 3 Figs. 13–14
Material: Six specimens in the Ko
¨seyahya section.
Remarks: This species is very rare in the Ko
¨seyahya
fauna. Most specimens have a smaller central area and less
bulbous and unequal arms. By its morphology, P. clavi-
formis is somehow intermediate between Paraparonaella
and Ropanella. Its tiny pores, rather smooth surface and the
absence of lateral distal projections are characters that
remind the latter genus, whereas the presence of an axial
spine at the distal end of each arm is a character of the
former genus. Unfortunately, the small number of speci-
mens and the poor preservation of most of them did not
permit us to section them to see if there is a microsphere
and, in affirmative case, how is its morphology.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Grossreifling, Austria; Ko
¨seyahya
section, SE Turkey.
Paraparonaella fragilis (Kozur & Mostler 1978)
(Fig. 6o, p)
1978 Rhopalodictyum fragilis Kozur & Mostler: 148, pl.
5 Fig. 4
1984 Paronaella fragilis (Kozur & Mostler).—Lahm:
89, pl. 16 Fig. 3
1999 Paronaella fragilis (Kozur & Mostler).—Tekin:
89, pl. 9 Fig. 8
Material: 13 specimens in 04-ELB 1 and 04-ELB 5.
Remarks: The arms of this rare species have the same
structure as those of the genus Triassoastrum Kozur & Mos-
tler, but the cortical layer is not so regular, forming polygonal
or oval meshes of various sizes. Some specimens also start
forming a thin spongy cover in the interradial areas.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Grossreifling, Austria; Bozkir, Konya,
and Ko
¨seyahya section, SE Turkey.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 367
123
Author's personal copy
Paraparonaella okuyucui Dumitrica & Tekin n. sp.
(Fig. 6q–u)
Description: Test with long, slender, approximately
equal and equally spaced arms, subcircular to subelliptical
in cross-section. Branches of primary beams lying in suc-
cessively inverted verticils of three. Cortical layer of the
arms with cortical beams more or less visible, composed of
large, polygonal pore frames with small nodes at pore
frame vertices. Tips of arms wide, wedge-shaped, usually
flattened, with much finer polygonal pore frames, and
usually three to five small, basally tricarinate, distally
needle-like spines of which a central one, aligned to the
primary beam, is usually distinguished.
Material: About 50 specimens in 04-ELB-1 and
04-ELB-2, of which ten have been photographed.
Holotype: Fig. 6q, Muse
´edeGe
´ologie Lausanne, No
MGL 96826
Dimensions: (in lm, based on ten specimens): Length of
arms 216–260, width of arms at the base 40–60, length of
distal spines 24–67.
Remarks: This new species differs from the holotype of
Paronaella simoni (Kozur and Mostler 1978) by having
longer and thinner arms with fewer distal spines and
wedge-shaped tips.
Etymology: The species is dedicated to Dr. Cengiz
Okuyucu, General Directorate of Mineral Research
and Exploration, Ankara, Turkey, in honor of his
contributions to the study of Paleozoic foraminiferal
biostratigraphy.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section,
eastern Turkey.
Paraparonaella parvispinosa (Kozur & Mostler 1978)
(Fig. 6v, w)
1978 Rhopalodictyum parvispinosum Kozur & Mostler:
149, pl. 5 Fig. 2
Description: Microsphere of tritrabid type. Arms equal
and equally spaced, broad, increasing in breadth distally.
Their tips rounded, usually with three thin spines, of which
one is axial and two are lateral, the former always longer
than the other two. Surface of test with irregular polygonal
pores frames. Arms with a central structure similar to that
of Triassoastrum surrounded by a thick spongy meshwork
developed especially on lateral sides. Spongy meshwork
not arranged in layers.
Material: Thirty specimens in the Ko
¨seyahya section.
Remarks: Morphologically, the species is very closed to
P. robusta (Kozur & Mostler), from which it differs in
having broader arms and rounded tips.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling, Austria; Ko
¨seyahya section,
eastern Turkey.
Paraparonaella robusta (Kozur & Mostler 1978)
(Fig. 6x)
1978 Rhopalodictyum robustum Kozur & Mostler: 150,
pl. 3 Fig. 18
1984 Paronaella robusta (Kozur & Mostler).—Lahm:
88, pl. 16 Fig. 2
Material: Fifteen specimens in the Ko
¨seyahya section.
Remarks: Morphologically, this species is very close to
Paraparonaella fragilis (Kozur & Mostler), from which it
differs by the presence of a thicker spongy cover around arms.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Grossreifling, Austria; Ko
¨seyahya
section, Elbistan, SE Turkey.
Genus Pseudangulobracchia Dumitrica & Tekin n. gen.
Type species:Pseudangulobracchia prisca n. sp.
Diagnosis: Tritrabids with quadrangular arms in cross-
section. Microsphere of tritrabid type, but with more or less
regular upper and lower faces. Arms with a primary beam
in the centre bearing verticils of three branches that form
three primary pores, and a spongy meshwork toward the
two faces. Verticils are successively inverted so that there
are no primary canals. Cortical shell without longitudinal
beams, with nodes interconnected by bars forming polyg-
onal (triangular to pentagonal) meshes.
Remarks: Although externally the test of this genus
resembles that of the middle Jurassic to Cretaceous genera
Angulobracchia Baumgartner and Halesium Pessagno, it
differs structurally from them by having a tritrabid instead
of angulobracchiid type of microsphere (see De Wever
et al. 2001). To this genus, we also assign the Rhaetian
species Paronaella pacofiensis Carter that seems to have
arms with the same rectangular cross section.
Etymology: From the Greek pseudo—false and angulo-
bracchia, a genus name. Feminine gender.
Range: Middle Carnian, Tetraporobrachia haeckeli Zone
of the Ko
¨seyahya section to upper Norian?-Rhaetian of
Kunga Group, Charlotte Islands, B.C. (Carter, 1993).
Pseudangulobracchia glenniei Dumitrica & Tekin n. sp.
(Fig. 7a, b)
368 P. Dumitrica et al.
123
Author's personal copy
Description: Test three-armed with arcuate interradial
space. Arms equal, thick, rectangular in cross section,
decreasing in breadth up to the inner third and then
increasing again distally and terminated in a club-shaped
tip. Cortical shell with wide rounded pores arranged rather
irregularly. Tips club-shaped with smaller pores and spin-
ules. Lateral sides of arms vertical with smaller pores than
upper and lower surfaces. Microsphere unknown but sup-
posed to be similar to that of type species.
Material: Two specimens in Glenn Shale, east-central
Alaska.
Holotype: Figure 7a, Muse
´edeGe
´ologie Lausanne, No.
MGL 96827.
Dimensions (in lm based on two specimens): Length of
arms 198–249, minimal breadth of arms 66–68, maximal
breadth of tips 132–149.
Remarks: The species differs from Pseudangulobrac-
chia prisca n. sp. and Pseudangulobracchia pacofiensis
(Carter) in having shorter and thicker arms and arcuate
interradial space.
Etymology: The species is named for W. K. Glennie, for
his contribution to the geology of the Oman Mountains.
Range and occurrence: Latest Tuvalian-earliest Lacian,
Glenn Shale, east-central Alaska, so far as known.
Pseudangulobracchia prisca Dumitrica & Tekin n. sp.
(Fig. 7c–f)
Description: Test with three equal narrow arms, which
are constant in width, except the distal ends. Arms qua-
drangular in cross-section, about twice higher than their
width. Edges of arms without longitudinal beams forming
keels. Tips widened distally, wedge shaped without spines.
They have a thinner spongy meshwork than that of arms,
which is usually radially arranged. Central area flat. Upper
and lower surfaces of test flat with small nodes disposed
disorderly or in short longitudinal or oblique rows and
interconnected by bars forming polygonal (triangular to
pentagonal) meshes. Lateral sides of arms straight, vertical.
Material: Tens of specimens in 04-ELB-2, of which five
have been photographed.
Holotype: Fig. 7c, Muse
´edeGe
´ologie Lausanne, No.
MGL 96828.
Dimensions (in lm, based on 4 specimens): Length of
arms 192–250, width of arms at base 28–43, width of tips
65–105.
Remarks: The species is very constant in its morphol-
ogy. By its general shape and rectangular to subrectangular
transverse section of arms, it resembles Paronaella
pacofiensis Carter from the late Norian?-Rhaetian of
Charlotte Islands that should probably be assigned to the
same genus.
Etymology: From the Latin priscus, -a,-um—very old,
ancient.
Range and occurrence: Middle Carnian late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section,
Elbistan, SE Turkey.
Genus Ropanaella Dumitrica & Tekin n. gen.
Type species:Ropanaella ungulata n. sp.
Diagnosis: Spongy three-armed spumellaria of unknown
microsphere. Central part of test small and low. Whole test
with a very dense and delicate spongy network, which is
more or less radially arranged on arms. No primary beam
visible inside arms. Tips not expanded, armed or not with
several short spines as external prolongations of internal
beams of the skeleton. Arms without axial spine or pylome
on tips. Surface usually smooth, without nodes or longi-
tudinal beams.
Remarks: This genus is erected for several late Ladinian
(Longobardian) and middle Carnian (late Julian) three armed
species with very delicate, dense spongy arms. Morpholog-
ically it resembles the genus Paronaella Pessagno, from
which it differs in having a dense spongy meshwork without
external ornamentation, in having no primary beam along
arms and no pylome. It differs from Paraparonaella n. g. in
having no axial spines as prolongation of primary beams, no
expanding tip, a rather smooth surface, and the spongy
meshwork made of very thin elements.
The microsphere of this genus is not known. All sections
we made in test have shown a very thin spongy meshwork,
mostly dissolved around a spherical hollow. This suggests
either that the microsphere is made of very delicate and
easily dissolved elements or that there is no microsphere.
Anyway, the microsphere, if there is one, is probably nei-
ther tritrabid nor angulobrachiid because of the absence of
primary beams along the central part of arms. Because of
all these reasons, the genus is just provisionally included
under the Tritrabidae.
Most species come from the middle Carnian, very few
species [Ropanaella glabra (Kozur & Mostler), R. spino-
folia (Kozur & Mostler) and Ropanella sp.], being known
to also occur in the middle-late Longobardian
Etymology: The name is an anagram of the genus name
Paronaella, which it morphologically resembles.
Range: Late Ladinian (Longobardian) Muelleritortis
cochleatata Zone, Spongoserrula rarauana Subzone to
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 369
123
Author's personal copy
Fig. 7 a,bPseudangulobracchia glenniei Dumitrica & Tekin n. sp.,
Glenn Shale: aholotype, bparatype, scale bar =200 lm. c–
fPseudangulobracchia prisca Dumitrica & Tekin n. sp.: cholotype,
04-ELB-2, scale bar =180 lm; d–fparatypes, all from 04-ELB-2,
edetail of the arm structure of Fig. d,scale bar =180, 55 and 180
lm, respectively. g-iRopanaella conobracchia Dumitrica & Tekin n.
sp.: gholotype, 04-ELB-5; h,iparatypes, 04-ELB-2, scale bar for all
specimens =200 lm. j–lRopanaella nitida Dumitrica & Tekin n.
sp.: jholotype, 04-ELB-2; k,lparatypes: k04-ELB-1, l04-ELB-5;
scale bar for all specimens =190 lm. mRopanaella spinofolia
(Kozur & Mostler), R78-6, scale bar =220 lm. n,oRopanaella
ungulata Dumitrica & Tekin n. sp.: nholotype 04-ELB-2, scale
bar =230 lm; oparatype, 04-ELB-2, scale bar =200lm. pRopan-
aella ygraeca Dumitrica & Tekin n. sp., holotype, 04-ELB-5, scale
bar =180 lm. q–tRopanaella zapfei (Kozur & Mostler): q04-ELB-
3, r04-ELB-4, s04-ELB-5, t04-ELB-2; scale bar for all
specimens =100 lm. uRopanaella sp., R78-6, scale bar =180
lm. v–xPlafkerium nazarovi Kozur & Mostler, Rc4, scale
bar =170, 240 and 140 lm, respectively
370 P. Dumitrica et al.
123
Author's personal copy
Middle Carnian Tetraporobrachia haeckeli Zone so far as
known.
Ropanaella conobracchia Dumitrica & Tekin n. sp.
(Fig. 7g–i)
Description: Arms relatively long, equal and equally
spaced, slightly conical, circular in cross-section, made of a
very delicate and dense spongy meshwork. Tips rounded
with numerous very small spines on distal end and also on
the lateral sides of tips. Surface of test smooth.
Material: Six illustrated specimens from samples
04-ELB-2 and 04-ELB-5.
Holotype: Fig. 7g, Muse
´edeGe
´ologie Lausanne, No.
MGL 96829.
Dimensions (in lm, based on six specimens): Length of
arms 246–308, diameter of arms at base 54–68, diameter of
arms at distal end 102–126.
Remarks: This species is close to R. ungulata n. sp. in
having the same type of dense test, small spines at tips, but
differs from it in having conical rather than flat arms.
Etymology: From the Latin conus—cone and bracchium,
-a—arm; a species with conical arms.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Ropanaella nitida Dumitrica & Tekin n. sp.
(Fig. 7j–l)
Description: Test small with a very compact spongy test
made of very thin meshwork. Arms smooth, quadrangular
to rounded-quadrangular in cross-section, increasing
slightly in thickness distally. Tips not expanded, slightly
rounded, without spines or with very small ones; some-
times the tip of one arm slightly forked. Sides vertical.
Material: Fifteen specimens in samples 04-ELB-1,
04-ELB-2 and 04-ELB-5.
Holotype: Fig. 7j, Muse
´edeGe
´ologie Lausanne, No.
MGL 96830.
Dimensions (in lm, based on six specimens): Length of
arms 208–290, width of arms at base 68–100, width of
arms at tip 110–149.
Remarks: This species differs from R. mocki (Kozur &
Mostler 1978) by its much denser spongy meshwork and
rectangular cross-section of arms.
Etymology: From the Latin nitidus, -a, -um - clean,
bright, elegant, etc.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, eastern Turkey.
Ropanaella spinofolia (Kozur & Mostler 2006)
(Fig. 7m)
2006 Paronaella spinofolia Kozur & Mostler: 26, pl. 7
Fig. 19
Remarks: The specimen herein illustrated and coming
from the Longobardian of the Rarau Mt., Eastern Carpa-
thians, very much resembles the holotype, from which it
only differs in having less broad arms and almost no lateral
spines, probably due to poor preservation.
Range and occurrence: Late Ladinian (middle to early
late Longobardian), Muelleritortis cochleata Zone, Spon-
goserrula rarauana and S. fluegeli Subzones; Dinarides
(Bosnia-Herzegovina) and Rarau Mountain, eastern Car-
pathians (Romania).
Ropanaella ungulata Dumitrica & Tekin n. sp.
(Fig. 7n-o)
Description: Test smooth with broad, moderately long
and approximately equally spaced arms consisting of very
fine and dense spongy meshwork. Arms elliptical in cross-
section, one of them slightly longer than the others and
slightly widened distally. Arms increasing in breadth dis-
tally, paw-shaped with one to five claw-like short spines.
Material: Two specimens.
Holotype: Fig. 7n, Muse
´edeGe
´ologie Lausanne, No.
MGL 96831.
Dimensions (in lm, based on two specimens): Length of
arms 222–244, breadth of arms at base 92–102, breadth of
arms at tips 153–168.
Remarks: This new species differs from Ropanaella
conobracchia n. sp. by having flat and broad rather than
conical arms, and claw-like tips. Although we have very
few specimens, this species has very characteristic features.
Etymology: From the Latin ungulatus, -a, -um—with
claws, with nails.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Ropanella ygraeca Dumitrica & Tekin n. sp.
(Fig. 7p)
Description: Test relatively long, Y-shaped with an axial
arm longer and slightly thicker than the other two and
separated from them by a wider angle. All arm cylindrical
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 371
123
Author's personal copy
or slightly decreasing in diameter distally. They are cir-
cular in cross-section, and made of a very delicate and
dense spongy meshwork. Surface of test smooth. Tips
transversally cut, armed with a few short and very thin
spines.
Material: A single specimen in the Ko
¨seyahya fauna,
sample 04-ELB-5.
Holotype: Fig. 7p, Muse
´edeGe
´ologie Lausanne, No.
MGL 96832.
Dimensions (in lm, based on holotype): Length of
longer arm 265, length of shorter arms 182, diameter of
arms 99.
Remarks: This species resembles R. conobracchiata n.
sp. and R. nitida n. sp. in having the same very fine spongy
test, but differs from both species in having cylindrical
rather than conical arms. From the former species, with
which it resembles most, R. ygraeca n. sp. differs in having
cylindrical arms rather than arms increasing in diameter
distally, in having no bulbous tips, and not being con-
stricted at the proximal end.
Etymology: From the Y-shaped of the test.
Range and occurrence: A single specimen in the
Ko
¨seyahya fauna, middle Carnian, Tetraporobrachia hae-
ckeli Zone, Ko
¨seyahya section, Elbistan, SE Turkey.
Ropanaella zapfei (Kozur & Mostler 1978) emended herein
(Fig. 7q–t)
1978 Dictyocoryne mocki Kozur & Mostler: 152, pl. 1
Fig. 7, non 9; pl. 5 Figs. 5, 6
1978 Dictyocorne zapfei Kozur & Mostler: 153, pl. 5
Figs. 7, 8
Emended description: Test very small, rough-surfaced,
consisting of a very delicate, dense meshwork. Arms short,
flattened, triangular in face view with thin, slightly serrate
distal ends. Central part small when simple, but thickened
when it is surrounded by a patagium-like or a spongy cover
that covers it on all sides, screening the connection of arms
at centre.
Material: Fifteen specimens.
Remarks: This species differs from the other species of
the genus Ropanaella in being rough-surfaced. Based on the
Ko
¨seyahya specimens, we consider that the two species (D.
mocki and D. zapfei) are synonyms, the former representing
specimens of the latter without patagium. We chose as name
of the species D. zapfei because it represents complete
specimens. The patagium is very well distinguished from
the arms by having a finer meshwork with smaller meshes.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling and Grossreifling, Austria;
Ko
¨seyahya section, Elbistan, SE Turkey.
Ropanella sp.
(Fig. 7u)
Remarks: It is probable that this specimen, coming from
the assemblage containing R. spinofolia (Kozur & Mos-
tler), belongs also to this latter species, but it is difficult to
assign to it because it has the lateral sides of arms corroded,
so that it is difficult to estimate the true breadth of its arms.
Range and occurrence: Late Ladinian (middle Longo-
bardian), Muelleritortis cochleata Zone, Spongoserrula
rarauana Subzone, Rarau Mt., Eastern Carpathians,
Romania.
2. Family VEGHICYCLIIDAE Kozur & Mostler 1972, emended
herein
Emended diagnosis: Spongy or spongy-like flattened
radiolarians consisting of a microsphere with commonly 12
pores that is at the origin of four coplanary or subcoplanary
primary beams. Beams may stop in the shell or may extend
outside it into spines or arms of hagiastrid structures.
Remarks: We include in this family all flat Triassc
spumellarian Radiolaria with four coplanar spines or
spongy arms, or only primary beams originated in a pen-
tagonal dodecahedral microsphere, which is one of the five
Platonic solids. This microsphere has a square or subsquare
equatorial ring with four primary beams in the equatorial
plane and, at the two poles, a pentagonal ring connected to
the square equatorial ring by five lateral bars (Fig. 8a, b).
The equatorial ring represents the zig-zagged line sepa-
rating the two halves of the pentagonal dodecahedron and
consists of ten bars. The presence of four equatorial pri-
mary beams in a dodecahedral polyhedron raises a problem
of symmetry, because on each side of the ring there are five
instead of four connecting bars between the square equa-
torial ring and the pentagonal polar ring. The solution that
these radiolarians found to solve this problem was the
following: from two opposite sides of the square arises one
connecting bar towards one pole and one towards the
opposite pole; and from the other two opposite sides, from
one side arise two bars towards one pole (let’s say upper)
and one bar (between the two bars) towards the other pole
(lower). At the opposite side, the situation is inverted; a
single bar arises towards the upper pole and two towards
the lower pole. The result is that the two halves (up and
down) of the microsphere are symmetrical only after a
rotation of 180°to one another. This inversion is the only
logical result of the dodecahedral shape of the microsphere
and the presence of only four primary beams.
372 P. Dumitrica et al.
123
Author's personal copy
In most of the studied specimens, the pentagonal
dodecahedron is rather imperfect (Fig. 8c–e, g, h); very
rarely, when the intervening bars are rather thick, it is
rather perfect (Fig. 8f). The imperfection of its geomet-
rical form is due to the delicacy of the constitutive bars
representing the edges of this solid. Most of these bars are
not straight, but curved or bent due to the bars arising
from them to form the second layer of test. Such bars of
second order do not arise from well-established places, as
is the case of the Hagiastridae, but seem to be randomly
chosen.
We do not yet perfectly know the volume of this group,
but it seems that it comprises not only Triassic, but also
some Jurassic and Early Cretaceous disc-shaped or lens-
shaped radiolarians (De Wever et al. 2001). The forerunner
of the whole group seems to be the genus Tetrapaurinella
Kozur & Mostler. It has all the features of an original
member: a) the oldest FAD (Pelsonian so far as known), a
non-differentiated spongy test, and spines with circular
cross section. All the other structural and morphological
characters such as: bladed spines, circular test with porous
equatorial plate, arms along primary beams, etc., are der-
ivated characters.
This microsphere differs essentially from the micro-
sphere and first system of the Jurassic and Cretaceous
Hagiastridae and Dactyliosphaeridae as described by
Fig. 8 Microspheres of the family Veghicycliidae. a–bFundamental
pentagonal dodecahedron outlined by these microspheres and the
position of the four primary beams (pr); alateral view showing
the position of the four primary beams and the zigzagged shape of the
equatorial ring; arrows indicate the side of the ring with two
connecting bars between it and the apical ring. bSame in axial or
polar view showing the pentagonal apical ring in the centre and the
equatorial ring around it. cmicrosphere, partly broken off, of
Triassocrucella goestlingensis (Kozur & Mostler); dmicrosphere,
partly broken off, of Triassocrucella baloghi (Kozur & Mostler);
emicrosphere of Plafkerium quadratum (Lahm); fmicrosphere of
Veghicyclia n. sp. from an early Tuvalian sample of Rhodos Island;
g,hmicrosphere of Praeorbiculiformella sp. Scale bar of 10 lm is for
Figures c–h
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 373
123
Author's personal copy
Dumitrica in De Wever et al. (2001, Figs. 76–78). In these
post-Triassic radiolarians, there is no dodecahedral
microsphere, and each polar structure is connected to the
square equatorial ring by only four bars. Because of this,
the two polar structures or cupolas are rotated to one
another around the polar axis only 90°, instead of 180°.
Also, the Triassic arm-bearing Veghicycliidae have no
systems of girdles and no pyloniacean growth as have the
Hagiastridae. So, despite the morphological similarity
between the arms of the arm-bearing Veghicycliidae and
the Hagiastridae, the former are not forerunners of the
latter.
Since we consider this type of microsphere as the pri-
mordial character of the family, we include in it several
subfamilies differentiated by the shape of test and the type
of projections along the primary beams, as follows:
subfamily Tetrapaurinellinae n. subfam., with lenticular
test and four spines as prolongations of the primary
beams,
subfamily Veghicycliinae Kozur & Mostler emend. herein,
with flat, circular test in the plane of the four primary
beams,
subfamily Triassocrucellinae n. subfam., with hagiastrid-
type arms around primary beams.
Range: Middle Anisian to probably Early Cretaceous (see
discussion under the range of the subfamily Veghicycliinae).
2a. Subfamily TETRAPAURINELLINAE Dumitrica & Tekin n.
subfam.
Type genus:Tetrapaurinella Kozur & Mostler 1994,
type species herein designated: Tetrapaurinella discoidalis
Kozur & Mostler 1994.
Diagnosis: Spumellarians with spherical or lenticular
spongy shell bearing four primary spines originated in a
microsphere with commonly 12 pores. Spines circular in
transverse section or three-bladed.
Remarks: Sections that one of us (PD) made in a Fass-
anian species of Tetrapaurinella and in other early Ladi-
nian or late Anisian species with 4 three-bladed spines and
lenticular spongy shell have proved that all have a pen-
tagonal dodecahedral microsphere similar to that of Veg-
hicyclia,Praeorbiculiformella and Triassocrucella, a fact
that proves close phylogenetical relationships between
them. Consequently, by its ancientness and less organized
test, we considered that this family is the forerunner of the
subfamilies Veghicycliinae and Triassocrucellinae. The
subfamily started with massive, non-bladed spines, and
continued with straight three-bladed spines, which very
soon started twisting around their axis.
Range: Middle Anisian (Pelsonian) to Rhaetian.
Genus Plafkerium Pessagno 1979, emended herein
Type species:Plafkerium abbotti Pessagno 1979.
1979 Plafkerium Pessagno.—Pessagno et al.: 179
1981 Paraplafkerium Kozur & Mostler: 72, type species
Paraplafkerium quinquespinosum Kozur & Mostler
1981:73
Emended diagnosis: Test spongy, lenticular with 4
three-bladed spines originated in a primitive hagiastrid
microsphere commonly with 12 pores.
Remarks: Since its publication, this genus was inter-
preted in different ways by different authors. Most authors
assigned to it species assignable to Muelleritortis Kozur or
to related genera (Kozur and Mostler 1981; Lahm 1984;
Gorican and Buser 1990, etc.). These different interpreta-
tions were due to the incomplete original diagnosis. Pess-
agno described this genus as having a cortical shell of two
layers, but he made no mention on the structure of a pos-
sible inner skeleton. There is no mention of a spongy test or
of an inner medullary shell. Apart from its description as
having a double-layered cortical shell, which is not at all
visible on the illustrated specimens, its resemblance with
the genus Muelleritortis was also due to the presence of a
spine longer than the others.
Although we do not think that we have definitely solved
the problem of this genus, that should start from the type
species, we think that it has a spongy test, as the test of the
species herein assigned to it. This possibility is based on
the test morphology, on the spongy aspect of its surface
with irregular nodes connected by bars, and on the presence
of twisted spines.
We also think that the genus Paraplafkerium Kozur &
Mostler is a junior synonym of Plafkerium. The holotype of
the type species of this genus is an anomaly, with an
additional smaller spine obliquely directed and originated
in an interradial area above the equatorial plane. Normally,
the species should have only four coplanar spines, as the
species Paraplafkerium alpinum Kozur & Mostler origi-
nally included in this genus. However, this species does not
belong to the genus Paraplafkerium. The number of its
spines and the double-layered structure of its cortical shell
prove that it is a species belonging to the family Hind-
eosphaeridae, and that it is very close to the species Pla-
fkerium ?firmum Gorican (in Gorican and Buser 1990) and
to the genus Muelleritortis Kozur. According to its
description and illustration, the type species of Para-
plafkerium has a spongy shell. Although we could not
study its internal structure, it should be similar to that of
Plafkerium nazarovi Kozur & Mostler, which we assign
herein to Plafkerium, and whose inner structure we now
374 P. Dumitrica et al.
123
Author's personal copy
know rather well. The earliest Ladinian or latest Anisian
species Tetraspongodiscus quadratus Lahm, a species that
has a spongy shell and a microsphere of veghicycliid type
should also belong to this genus.
The genus Plafkerium, as herein emended, differs from
Tertrapaurinella in having three-bladed spines and from
Tetraspongodiscus Kozur & Mostler in having spines
without lateral spinules. Thin sections made in specimens
of the species Plafkerium ?nazarovi Kozur & Mostler
prove that the microsphere of this genus is similar to that of
the type genus of the subfamily and of the other genera
included in it.
Range: Latest Anisian or earliest Ladinian to Rhaetian.
Plafkerium nazarovi Kozur & Mostler 1981
(Fig. 7v–x, 10a, b)
1981 Plafkerium ?nazarovi Kozur & Mostler: 72, pl. 57
Fig. 1, pl. 58 Fig. 1
1984 Tetraspongodiscus nazarovi (Kozur & Mostler).—
Lahm: 59, pl. 10 Fig. 7
1999 Tetraspongodiscus nazarovi (Kozur & Mostler).—
Tekin: 122, pl. 22 Fig. 10
Description: Test spongy, disc-shaped, and square to
practically circular in face view. Spongy shell not organized
in concentric layers. Spines thin and long, twisted dextrally
up to 360°around their axes, especially in their middle
portion, and terminated in a long needle-shaped spike.
Remarks: Of all the specimens assigned to this species,
only two differ significantly from the holotype and para-
type. One, not illustrated, has the spines twisted twice,
whereas practically the majority of spines are, at most, only
twisted once (360°). The other one (Fig. 7x) has a very
small central test, and because of this, it is herein considered
as being a young ontogenetic stage. It is interesting that this
species remained practically unchanged from the latest
Anisian or earliest Ladinian up to the middle Carnian.
Range and occurrence: Latest Anisian or earliest Ladi-
nian to middle Carnian: Recoaro, Italy; Grossreifling,
Austria; Bozkir, Konya, Ko
¨seyahya section, Elbistan, SE
Turkey.
Plafkerium odoghertyi Dumitrica & Tekin n. sp.
(Fig. 10c, d)
Description: Shell small, spongy, lens-shaped, and
squarish to subcircular in face view. Surface rough with
small, irregular and irregularly arranged pores. Spines poin-
ted, 1.5 to twice as long as diameter of test, sinistrally twisted
about 360°, and decreasing gradually in diameter distally.
Material: Three specimens, of which two are in the
Ko
¨seyahya section (04-ELB-2 and 04-ELB-5) and one in
the Glenn Shale, Alaska.
Holotype: Fig. 10c, Muse
´edeGe
´ologie Lausanne, No.
MGL 96833.
Dimensions (in lm based on two specimens): Length of
sides of shell 74–91, length of spines 116–140.
Remarks: The specimen resembles very much P. (?)
sinistrum n. sp. from which it differs in having spines
which are less twisted (less than 360°) and gradually and
equally decreasing distally in diameter.
Etymology: The species is named for Dr. Luis
O’Dogherty, Spain, to honour his important contributions
to the knowledge of Mesozoic radiolarians.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE Tur-
key; latest Carnian or early Norian, Glenn Shale, Alaska.
Plafkerium patrulii Dumitrica & Tekin n. sp.
(Fig. 10e, f)
Description: Test squarish, flat with straight sides and
slightly inflated faces. Surface of test nodose with nodes
interconnected by thin bars resulting in a reticulate aspect
with triangular or quadrangular meshes. Spines equal with
or shorter than diameter of test. They are three-bladed and
strongly (about 360°) dextrally twisted. Distally they
decrease quickly in diameter and terminate in a spike.
Material: The two illustrated specimens in sample R78-
6.
Holotype: Fig. 10e, Muse
´edeGe
´ologie Lausanne, No.
MGL 96834.
Dimensions (in lm based on two specimens) Length of
sides of spongy test 121–135, length of spines 97–145.
Remarks:P. patrulii n. sp. differs from P. nazarovi
Kozur & Mostler in having shorter and strongly twisted
spines, and test surface with robust nodes. Its spines and
test surface resemble very well those of the co-occurring
species Angulopaurinella carpatica n. sp.
Etymology: The species is named for the Romanian
geologist Dan Patrulius (1921–1982) for his great contri-
butions to the geology and paleontology of the Mesozoic of
Romanian Carpathians.
Range and occurrence: Late Ladinian (middle Longo-
bardian), Muelleritortis cochleata Zone, Spongoserrula
rarauana Subzone, Zimbru Series, Rarau Mt., Eastern
Carpathians.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 375
123
Author's personal copy
Plafkerium quadratum (Lahm 1984)
(Figs. 8e, 9a,10g–h)
1984 Tetraspongodiscus quadratus Lahm: 59, pl. 10 Fig. 6
Description: Test flat, rounded square to square with
small irregular pores. Spines three-bladed, with parallel or
subparallel side proximally and pointed distally, straight or
slightly dextrally twisted.
Remarks: Of all the specimens we included in this spe-
cies, a single specimen (Fig. 10h) differs significantly from
the holotype. It has the same type of spines, but its shell has
slightly concave sides. This feature resulted probably by
the extension of the spongy shell on the base of the spines
in mature ontogenetic stages.
Range and occurrences: Latest Anisian or early Ladi-
nian, Buchenstein Formation, Recoaro, North Italy.
Fig. 9 Equatorial sections through some species of the family
Veghicycliidae. aPlafkerium quadratum (Lahm), lower Ladinian,
Rc4, scale bar =42 lm. bsame, much magnified, showing the
microsphere, scale bar =21 lm. cTriassocrucella goestlingensis
(Kozur & Mostler), 04-ELB 1, scale bar =42 lm. dPraeorbiculi-
formella polyspinosa Kozur & Mostler, detail of central shell with
microsphere, same as Fig. g,scale bar =21 lm. eTriassocrucella
kamatai Dumitrica & Tekin n. sp., paratype, 04-ELB-2, scale
bar =42 lm. fVeghicyclia austriaca Kozur & Mostler, detail
showing the four cross-wisely disposed primary beams, 04-ELB-1,
scale bar =37 lm. gPraeorbiculiformella polyspinosa Kozur &
Mostler, same specimen as Fig. d, showing the 4 cross-wisely
disposed primary beams, 04-ELB-2, scale bar =46 lm
376 P. Dumitrica et al.
123
Author's personal copy
Fig. 10 a,bPlafkerium nazarovi Kozur & Mostler, 04-ELB-1, scale
bar =260 lm. c,dPlafkerium odoghertyi Dumitrica & Tekin n. sp.:
cholotype, 04-ELB-2, scale bar =120 lm; dparatype, 04-ELB-5,
scale bar =120 lm. e,fPlafkerium patrulii Dumitrica & Tekin n.
sp.: eholotype, R78-6, scale bar =140 lm; fparatype, R78-6, scale
bar =140 lm. g,hPlafkerium quadratum (Lahm), Rc4, scale
bar =150 lm. i,jPlafkerium ?sinistrum Dumitrica & Tekin n. sp.:
iholotype, 04-ELB-1, scale bar =100 lm; jparatype, 04-ELB-5,
scale bar =100 lm. kPlafkerium yehae Dumitrica & Tekin n. sp.:
holotype, 04-ELB-5, scale bar =75 lm. lPlafkerium sp. A, Rc4,
scale bar =100 lm. m,nTetraspongodiscus bipolaris Dumitrica &
Tekin n. sp.: mholotype, 04-ELB-2, scale bar =130 lm; nparatype,
04-ELB-2, scale bar =100 lm. o,pTetraspongodiscus longispinosus
Kozur & Mostler, 04-ELB-2, scale bar =120 lm. q–sVeghicyclia
austriaca Kozur & Mostler: q04-ELB-4, r04-ELB-1, s04-ELB-2;
scale bar for all specimens =160lm. t–vVeghicyclia centrodepressa
Dumitrica & Tekin n. sp: t,uholotype, 04-ELB-4; vparatype,
04-ELB-4; scale bar for both specimens =190 lm. w,xVeghicyclia
multispinosa Kozur & Mostler, 04-ELB-2, scale bar =200 lm
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 377
123
Author's personal copy
Plafkerium ?sinistrum Dumitrica & Tekin n. sp.
(Fig. 10i, j)
Description: Central shell very small, subspherical and
spongy, arranged in two to four perimicrospheric layers.
Microsphere square in face view, each cupola seemingly
connected to the equatorial ring by four bars. First two
layers square in face view with wide polygonal meshes.
Following layer subglobular with polygonal to hexagonal
pores. Last layer, when present, very delicate. Spines long,
coplanar, three-bladed, and once to twice sinistrally twis-
ted. They may start twisting from their base or a little later.
Their diameter is usually equal on the proximal half and
decreases slowly to the pointed distal end.
Material: Five specimens in samples 04-ELB-1 and
04-ELB-5, of which three were photographed.
Holotype: Fig. 10i, Muse
´edeGe
´ologie Lausanne, No
MGL 96835
Dimensions (in lm based on three specimens): Diameter
of central shell 71–81, diameter of spines at base 21–29,
length of spines 107–145.
Remarks: This species is questionably assigned to the
genus Plafkerium because its microsphere seems to be
simpler, with only eight or nine pores instead of 12.
Unfortunately, the limited number of specimens and their
preservation did not allow sectioning more than one specimen,
and the preservation of its microsphere is not perfect. From
P. nazarovi Kozur & Mostler this new species differs not only
in having a probably simpler microsphere, but also in having a
smaller shell and sinistrally twisted spines. The Longobardian
species Plafkerium quinquespinosum (Kozur & Mostler),
from which it especially differs in having a smaller test, has the
same type of spines.
Etymology: From the Latin sinistral, -tra, -trum—left,
anticlockwise, due to its sinistral twisting.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Plafkerium yehae Dumitrica & Tekin n. sp.
(Fig. 10k)
Description: Test spongy, square and flat with 4 three-
bladed spines at corners. Pores of test irregular, rounded.
Spines relatively short and thick, with thin and broad
blades. Blades increase in breadth distally up to the second
third, then descrease fast up to the pointed distal end.
Spines twisted sinistrally up to 90°or more. Twisting is
slower proximally and faster on the middle portion.
Material: A single specimen in sample 04-ELB-5.
Holotype: Fig. 10k, Muse
´edeGe
´ologie Lausanne, No.
MGL 96836.
Dimensions (in lm based on holotype): Length of sides
of test 37, length of spines 77.
Remarks: Remains of test at the base of spines suggest
that the spongy test of the holotype is incomplete, the
outermost layer not being built yet or being dissolved
during fossilization. Although we have a single speci-
men of this type, by its morphology it differs rather
significantly from all the species of the genus so far
described.
Etymology: The species is named for Dr. Kuei-Yu Yeh,
National Museum of Natural Science, Taichung, Taiwan,
to honour her contributions to the knowledge of Mesozoic
radiolarians.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Eastern, SE
Turkey.
Plafkerium sp. A
(Fig. 10l)
Remarks: This species, of which we have only the
illustrated specimen, has a shell similar to that of P.
quadratum Lahm, but its spines are slightly sinistrally
twisted, and two of them are thicker.
Range and occurrences: Early Ladinian, Buchenstein
Formation, Recoaro, North Italy.
Genus Tetraspongodiscus Kozur & Mostler 1979, emended
herein
Type species:Tetraspongodiscus longispinosus Kozur &
Mostler 1979.
Emended description: Microsphere as with the sub-
family. Test spongy, lenticular with four equatorial three-
bladed spines. Spines bearing a verticil of spinules. Smaller
polar spines possibly present.
Remarks: We restrict the diagnosis of the genus to the
species with spongy shell and spines bearing a verticil of
spinules. The species with simple three-bladed spines are
assigned to the genus Plafkerium (see above). Kozur &
Mostler (1979) included also in this genus Staurodoras
dercourti De Wever, a taxon that contains two species:
the holotype is a species of the genus Pentaspongodis-
cus (see Lahm 1984), whereas the paratype should
be probably assigned to Kahlerosphaera Kozur &
Mostler.
378 P. Dumitrica et al.
123
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Range: Middle Carnian, Tetraporobrachia haeckeli
Zone.
Tetraspongodiscus bipolaris Dumitrica & Tekin n. sp.
(Fig. 10m, n)
Description: Test globular, spongy with four spines in
equatorial plane and two smaller in the polar axis. Micro-
sphere as with the family. Equatorial spines originated in
the corners of the microsphere, polar spines seem to be
originated in one side of the upper and lower pentagons.
Spongy shell not organized in concentric layers and rather
loose. Cortical shell with numerous and very small pores.
Polar spines thin, three-bladed, and pyramidal. Equatorial
spines thick, three-bladed, with parallel sides, each blade
bearing a laterally directed spinule. Spinules straight or
slightly recurved. Beyond verticils, distal ends of equato-
rial spines are low pyramidal or slightly rounded.
Material: Six specimens in sample 04-ELB 2, of which
two were photographed.
Holotype: Fig. 10m, Muse
´edeGe
´ologie Lausanne, No.
MGL 96837.
Dimensions (in lm based on three specimens): Diameter
of spongy shell 66–90, length of polar spines 50–79, length
of equatorial spines 72–109.
Remarks: The test of this new species very much
resembles that of Tetraspongodiscus longispinosus Kozur
& Mostler, from which it only differs in having two polar
spines and a globular shell.
Etymology: From having two poles.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Tetraspongodiscus longispinosus Kozur & Mostler 1979
(Fig. 10o, p)
1979 Tetraspongodiscus longispinosus Kozur &
Mostler: 81, pl. 11 Fig. 1
non 1984 Tetraspongodiscus longispinosus Kozur &
Mostler.—Lahm: 60, pl. 10 Fig. 9
Remarks: Our specimens differ from the holotype in
having slightly longer and recurved spinules and the cor-
tical layer of the spongy shell corroded. The specimen
included by Lahm (1984) in this species has a latticed
cortical shell and belongs to another genus.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling, Austria; Ko
¨seyahya section,
Elbistan area, eastern Turkey.
2b. Subfamily VEGHICYCLIINAE Kozur & Mostler 1972,
emended herein
Type genus:Veghicyclia Kozur & Mostler 1972
1972 Veghicyclidae Kozur & Mostler: 9
2006 Spinohollisellidae Kozur & Mostler: 39
Emended diagnosis: Disc-shaped or lens-shaped spum-
ellarians with four primary beams originated in the corners
of a simple, delicate microsphere, commonly with 12
pores. Primary beams originated in the middle of some,
bars not in triclad junctions. Extramicrospheric test latticed
or spongy, flat, and circular.
Remarks: Kozur & Mostler (2006) considered that the
microsphere of this subfamily is imperforate, having only
nodes as in many Late Paleozoic Ruzhencevispongacea,
but they produced no proofs supporting this affirmation. On
the contrary, our investigations under optical and electron
microscopes, based on the very well preserved specimens
from the Ko
¨seyahya samples and from other samples
(Figs. 8,11b, d, g, h, 12d), show that the microsphere
consists of very thin bars and that it usually has 12 wide
polygonal pores and a very primitive structure of hagiastrid
type. Based on this structure, to which we give a primary
value at the family and subfamily levels, we emended the
diagnosis of the subfamily, including in it not only the
genus Praeorbiculiformella Kozur & Mostler, but also
the family Spinohollisellidae, represented by the nominal
genus (Spinohollisella Kozur & Mostler 2006) and proba-
bly a single species, although originally four species have
been described (Kozur & Mostler 2006). The microsphere,
and in fact the inner structure of this genus is not yet
known, but we suppose that it could be similar to that of the
other genera included in the Veghicycliidae. The differ-
ences that Kozur & Mostler (2006) mentioned to motivate
the description of this new family have, in our opinion, just
a generic value.
We reject from this subfamily the genera Palaeolitho-
cyclia Kozur & Mostler, Palaeoastrocyclia Kozur &
Mostler, Ruesticyclia Kozur & Mostler, and Praetrigono-
cyclia Kozur & Mostler, erected by Kozur and Mostler
(1972) on the basis of the species described and illustrated
from thin sections by Ru
¨st (1885,1892,1898). These taxa
have generally no taxonomic value and should be consid-
ered nomina dubia, because of the difficulty or impossi-
bility of determining their true status and of recognizing them
in the fossil material. Erection of such genera does not
advance our knowledge of the taxonomy and evolution of this
group but, on the contrary, produces confusions because it
burdens the taxonomy with useless names.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 379
123
Author's personal copy
The subfamily Veghicycliinae is not a member of the
family Saturnalidae as Bragin (2007) considered, because
of its morphology and especially because of the initial
structure of the skeleton of its members. The Saturnalidae
generally have a peripheral ring and a heteropolar micro-
sphere with a special structure, which is at the origin of two
polar arms (Dumitrica 1985, Dumitrica and Zu
¨gel 2008,De
Wever et al. 2001), whereas the Veghicycliinae have
another type of microsphere, four primary beams and never
a saturnalid ring.
The subfamily Veghicyclinae seems to have appeared
from the subfamily Intermediellinae, and especially from
the genus Tetrapaurinella, by the development of an
equatorial plate and shortening of the four spines (genus
Veghicyclia) and/or development of a spongy skeleton at
the periphery of the spongy lenticular shell (genus Prae-
orbiculiformella). One of the transitional species between
the latter genus and Tetrapaurinella could be Orbiculifor-
ma gazipasaensis Tekin with its peripheral spongy ring.
Range: Late Ladinian (Longobardian) to Rhaetian (see
Baumgartner et al. 2008), or even to Early Cretaceous
(Hauterivian) so far as known. The extended range of the
family up to the Hauterivian is based on the presence of
several undescribed species with an equatorial plate similar
to that of Veghicyclia in the Middle Jurassic (Bajocian)
(Spongotrochus sp. B3 in Nishihara, 2009), Late Jurassic
(Oxfordian and Early Tithonian undescribed species in the
senior coauthors’s collection), and Early Cretaceous (one
species illustrated as Pseudoaulophacus ?florealis Jud in
Baumgartner et al. 1995, p. 439, pl. 5334 Fig. 4; and as
unnamed veghicycliid in De Wever et al. 2001, Fig. 75.2,
and another species illustrated as Sciadiocapsa (?) sp. A in
Ishii et al. (2009, pl. 16 Fig. 2). Although the microsphere
of these younger species is probably not perfectly similar to
that of the Veghicycliidae, the presence of four coplanar
primary beams, of a perforate equatorial plate, and the
general morphology of their spongy test are elements in
common with the Triassic taxa so that, for the moment, it is
impossible to classify them in another family group. To
assign them, even questionably, to Sciadiocapsa is
impossible because this genus is a nassellarian and not a
spumellarian.
Genus Veghicyclia Kozur & Mostler 1972, emended herein
Type species:Veghicyclia pulchra Kozur & Mostler
1972.
1972 Veghicyclia Kozur & Mostler: 10
Emended diagnosis: Veghicycliidae with an equatorial
latticed circular plate covered on both sides by a spongy
test of various sizes, but usually reduced to the central part
around the microsphere. Pores of equatorial plate radially
or irregularly arranged. Border of plate bearing usually a
variable number of spines.
Range: Middle Carnian to Rhaetian or younger.
Veghicyclia austriaca Kozur & Mostler 1972
(Figs. 9f, 10q–s)
1972 Veghicyclia austriaca Kozur & Mostler: 12, pl. 3
Fig. 3; pl. 4 Figs. 10, 15
1972 Veghicyclia robusta Kozur & Mostler: 15, pl. 3
Figs. 1, ?4, 7
1984 Veghicyclia austriaca Kozur & Mostler.—Lahm:
94, pl. 17 Fig. 6
1999 Veghicyclia austriaca Kozur & Mostler.—Tekin:
117, pl. 20 Fig. 1
Material: More than 100 specimens.
Remarks: In agreement with Tekin (1999), we consider
that V. robusta Kozur & Mostler is a synonym of V. aus-
triaca Kozur & Mostler. Both ‘‘species’’ occur in the
Ko
¨seyahya samples and have the same type of radial and
circular arrangements of pores.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling and Grossreifling, Austria;
Bokzir, Konya and Ko
¨seyahya section, Turkey.
Veghicyclia centrodepressa Dumitrica & Tekin n. sp.
Fig. 10t–v
Description: Equatorial plate circular, with numerous,
disorderly arranged and very small pores, and more than 15
peripheral spines. On both faces, middle and central parts
of equatorial plate covered with a spongy test that has a
raised circular area at about the middle of radius and a wide
central depression. Slope of raised area gentle both towards
periphery and centre of test. Central depression with a
small raised area in the centre. Pores of spongy test small
on the raised circular area and on the external slope, and
very fine in the central depression. Peripheral spines short,
triangular and irregularly spaced.
Material: Two specimens in ample 04-ELB-4 .
Holotype: Fig. 10t, u, Muse
´edeGe
´ologie Lausanne, No.
MGL 96838.
Dimensions (in lm, based on two specimens): Diameter
of shell without spines 244–341, length of spines 68–78.
Remarks: Although only based on two specimens, this
species is very well characterized and distinguished from
all the species so far known by having a raised circular area
380 P. Dumitrica et al.
123
Author's personal copy
and a central depression. By these characters, this species
could be also assigned to the genus Praeorbiculiformella
Kozur & Mostler, but it differs from the species of this
genus in having a perforate equatorial plate.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Veghicyclia multispinosa Kozur & Mostler, 1972
(Figs. 10w, x, 11a, b)
1972 Veghicyclia multispinosa Kozur & Mostler: 13, pl.
3 Fig. 5
1984 Veghicyclia multispinosa Kozur & Mostler.—
Lahm: 96, pl. 17 Fig. 10
Material: About 50 specimens.
Remarks: The difference between this species and V.
reiflingensis Kozur & Mostler is discussed under the latter
species.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone: Go
¨stling and Grossrei-
fling, Austria; Ko
¨seyahya section, Elbistan, SE Turkey.
Veghicyclia pauciperforata Kozur & Mostler 1972
(Fig. 11c–e)
1972 Veghicyclia pauciperforata Kozur & Mostler: 14,
pl. 3 Figs. 8, 12
1984 Veghicyclia pauciperforata Kozur & Mostler.—
Lahm: 96, pl. 17 Fig. 9
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone: Grossreifling, Austria;
Ko
¨seyahya section, Elbistan, SE Turkey.
Veghicyclia radiata Dumitrica & Tekin n. sp.
(Fig. 11f–k)
1972 ?Veghicyclia robusta Kozur & Mostler, part: 14, pl.
3 Fig. 4, non 1, 7
Description: Test represented especially by the equato-
rial plate with a very small globular center comprising the
microsphere and a thin spongy cover. Equatorial plate with
a very well visible radial structure commonly consisting of
12–14 rays, all extended outside into spines. Four rays are
disposed crosswise, start from the corners of the micro-
sphere and represent the primary beams. The other rays are
interradial and start from the perimicrospheric spongy
meshwork. In each interradial interval there are two,
maximum three such rays. Outside all peripheral spines are
of relatively similar length. Pores increase in size distally
and are disposed both radially and more or less concen-
trically. Since pores are also radially disposed and each
interradial interval has only one pore, rarely two, in the
tangential direction, pores become larger and larger distally
and tangentially elongated.
Material: More than 30 specimens in samples 04-ELB-1
and 04-ELB-2.
Holotype: Fig. 11f–h, Muse
´edeGe
´ologie Lausanne, No.
MGL 96839, .
Dimensions: (in lm, based on eight specimens): Diam-
eter of shell without spines 194–274, length of spines
25–62.
Remarks: This is the most frequent species of Veghi-
cyclia in the Ko
¨seyahya fauna.
Etymology: From its radial structure.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone: Ko
¨seyahya section, Elb-
istan, SE Turkey; probably also in Grossreifling, Austria.
Veghicyclia reiflingensis Kozur & Mostler 1972
(Fig. 11l–n)
1972 Veghicyclia reiflingensis Kozur & Mostler: 14, pl.
3, Fig. 6
1984 Veghicyclia reiflingensis Kozur & Mostler.—
Lahm: 95, pl. 17 Fig. 8
Description: Skeleton consisting of a large equatorial
latticed plate and a very small spongy spherical central part
around the microsphere. Equatorial plate thin, with wide,
irregularly arranged polygonal pores and thin intervening
bars. Periphery of plate irregular, ragged, usually with
25–30 thin spines of various length and arrangement. The
four primary beams well marked across the plate and
prolonged into spines beyond it.
Material: More than 40 specimens.
Remarks: The specimen that shows the best the mor-
phology of this species is the topotype specimen illustrated
by Lahm (1984) from Grossreifling. The great number of
specimens we found in the Ko
¨seyahya samples shows that
between this species and V. multispinosa Kozur & Mostler
there are many transitional forms, but that most specimens
of each species are morphologically close to their holo-
types. What differentiates this species from V. multispinosa
are: a finer equatorial plate, larger pores, an irregular
periphery as if the plate is not finished, and, usually, the
absence of a well-marked poreless peripheral band.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 381
123
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Fig. 11 a,bVeghicyclia multispinosa Kozur & Mostler, 04-ELB-2:
bdetail of the central part of Fig. a,scale bar =200 and 40 lm,
respectively. c–eVeghicyclia pauciperforata Kozur & Mostler,
04-ELB-1: ddetail of the central part of Fig. cwith part of
microsphere broken off; scale bar =170, 20 and 170 lm, respec-
tively. f–kVeghicyclia radiata Dumitrica & Tekin n. sp.: f–hholotype,
04-ELB-2, gand hdetail of the microsphere, scale bar =150, 15 and
15 lm, respectively; i–kparatypes, i04-ELB-2, j,k04-ELB-1, scale
bar for all specimens =150 lm. l–nVeghicyclia reiflingensis Kozur
& Mostler, 04-ELB-2, scale bar for all specimens =180 lm.
o,pVeghicyclia sp. A: o04-ELB-1, p04-ELB-5, scale bar for both
specimens =110 lm. qVeghicyclia sp. B, 04-ELB-1, scale
bar =110 lm. rVeghicyclia sp. C, 04-ELB-1, scale bar =110
lm. s–uCarinacyclia acari Dumitrica & Tekin n. sp.: sholotype,
04-ELB-2, scale bar =130 lm; t,uparatypes, t04-ELB-2,
u04-ELB-5, scale bar for both Figures =130 lm. vCarinacyclia
sol Dumitrica & Tekin n. sp.: holotype, 04-ELB-2, scale bar =230
lm. w,xPraeorbiculiformella alaskaensis Dumitrica & Tekin n. sp.:
wholotype, Glenn Shale, scale bar =200 lm; xparatype, Glenn
Shale, scale bar =200lm
382 P. Dumitrica et al.
123
Author's personal copy
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone: Grossreifling, Austria;
Ko
¨seyahya section, Elbistan, SE Turkey.
Veghicyclia sp. A
Fig. 11o, p.
Remarks: In this species we include the two illustrated
specimens, the only ones found so far, characterized by an
irregular equatorial plate with a variable number of small
peripheral spines, small and irregularly arranged pores, and
a small spongy central shell.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section, Elb-
istan, SE Turkey.
Veghicyclia sp. B
Fig. 11q
Remarks: This morphotype, from which we have only
the illustrated specimen, looks like an anomalous speci-
men, having an irregularly undulate equatorial plate with
oval pores that are radially oriented.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section, Elb-
istan, SE Turkey.
Veghicyclia sp. C
Fig. 11r
Remarks: This incompletely preserved and small speci-
men, with periphery without spines and a central spongy
test, looks like an anomalous specimen.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section, Elb-
istan, SE Turkey.
Genus Carinacyclia Kozur & Mostler 1972, emended
herein
Type species:Carinacyclia costata Kozur & Mostler
1972.
Emended definition: Flat skeleton consisting of a porous
equatorial plate covered on both sides by a porous plate and
bearing peripheral spines. Initial skeleton consisting of a
microsphere, commonly with 12 pores and four primary
beams running across the equatorial plate.
Remarks: The genus, initially described on the basis of
the type species, was considered as havinga spongy to porous
test. The study of the species Carinacyclia acari n. sp. in
optical microscopy proves undoubtedly that the skeleton is not
spongy, but consists of two porous plates that cover the
equatorial plate on both sides. The microsphere and number of
primary radial beams are similar to those of Veghicyclia and
Praeorbiculiformella. A small, delicate spongy shell exists
only around the microsphere, as in the other two genera.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling, Austria; Ko
¨seyahya, eastern
Turkey.
Carinacyclia acari Dumitrica & Tekin n. sp.
Fig. 11s–u
Description: Equatorial plate covered on both sides by a
flat porous plate, which can extend up to the border of the
plate. Both these plates and equatorial plate with rounded
polygonal pores of various size and irregular arrangement.
Centre of test flat or with a very small inflation consisting of a
very delicate spongy meshwork around the microsphere.
Margin of both faces rounded, without a keel. Margin of plate
with 12–27 triangular spines, which are usually relatively
short but their length differs especially with specimen.
Material: More than 50 specimens in Ko
¨seyahya section
samples.
Holotype: Fig. 11s, Muse
´edeGe
´ologie Lausanne, No.
MGL 96840.
Dimensions (in lm, based on 17 specimens): Diameter
of shell without spines 218–266, length of spines 27–50.
Remarks: By its shape, pore size, and type and number
of spines this species resembles Carinacyclia costata Ko-
zur & Mostler, from which it differs in having no keel on
the periphery of the spongy test. The variability of the
species is especially expressed in the number of peripheral
spines and diameter of shell.
Etymology: The species is named for Sukru Acar
(General Directorate of Mineral Research and Exploration
(MTA), Ankara, Turkey) in honour of his contributions to
the biostratigraphy of Tertiary Foraminifera.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section, Elb-
istan, SE Turkey.
Carinacyclia sol Dumitrica & Tekin n. sp.
(Fig. 11v)
Description: Test circular, flat, with equatorial plate
covered on both sides by a porous plate, the diameter of
which is slightly smaller than the diameter of equatorial
plate. Pores of this plate small, polygonally framed,
unequal, and irregularly arranged. Border of equatorial
plate outside the cortical plates very narrow and armed
with some 30 peripheral spines, which are relatively short,
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 383
123
Author's personal copy
thin, and disposed at rather regular distances from one
another.
Material: A single specimen in the Ko
¨seyahya samples.
Holotype: Fig. 11v, Muse
´edeGe
´ologie Lausanne, No.
MGL 96841.
Dimensions (in lm based on one specimen): Diameter of
test without spines 452, length of spines 81.
Remarks: Although we have a single specimen, by the
number of its peripheral spines (28), which is almost
double by comparison to that of C. acari n. sp., their small
size, and the greater diameter, this specimen is too different
from the other species of this genus and can very easily be
considered as representing a new species.
Etymology: From the Latin sol—sun.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone, Ko
¨seyahya section, Elb-
istan, SE Turkey.
Genus Praeorbiculiformella Kozur & Mostler 1978
Type species:Praeorbiculiformella plana Kozur &
Mostler 1978
1978 Praeorbiculiformella Kozur & Mostler: 163
Remarks:Praeorbiculiformella differs from Veghicyclia
in that its skeleton is thicker on the periphery than in the
centre, and in having the pores arranged in more or less
regular concentric circles. The range of the genus was
recently considered early Carnian—late Carnian (O’Dogh-
erty et al. 2009), but it could be extended to late Ladinian, if
we include in it Orbiculiforma gazipasaensis Tekin, and up
to Rhaetian, if we include in it Orbiculiforma multibrachiata
Carter, translated to Orbiculiformella by Baumgartner et al.
(2008). Since the type species of Orbiculiformella Kozur &
Mostler (O. railensis Pessagno) is a mid-Cretaceous (late
Albian) species, and the genera Orbiculiforma and Orbicu-
liformella seems to be polyphyletic, we think that Carter’s
species can be considered to be either one of the last mem-
bers of Praeorbiculiformella or a type species of a new
genus that would differ from this Triassic genus by having
spongy peripheral arms instead of spines.
Range: Late Ladinian to early Norian or Rhaetian.
Praeorbiculiformella alaskaensis Dumitrica & Tekin n. sp.
(Fig. 11w, x)
Description: Test circular consisting of an equatorial
plate and a wide rim. Centre not known because of poor
preservation, but supposed to be small and raised as in most
species. Central cavity as wide as half the ray of test, very
thin consisting of the equatorial plate with circular or
subcircular pores disposed in rather disturbed concentric
rows and decreasing in size toward the centre. Rim thick
and wide with sharp, vertical inner boundary and gentle
slope toward the periphery. It has one inner row of rounded
subrectangular pores and one or two incomplete and
irregular rows on the middle and outer side. All pore
frames have small nodes at vertices. Periphery with a
porous keel representing prolongation of the equatorial
plate. It has an incomplete row of pores and about 20 flat,
lamellar triangular spines or numerous small and flat
peripheral spines representing intervening bars of an
incompletely built peripheral row of pores.
Material: Four specimens in the Glenn Shale, east-cen-
tral Alaska, of which only three have been photographed.
Holotype: Fig. 11w, Muse
´edeGe
´ologie Lausanne, No.
MGL 96841.
Dimensions (in lm based on three specimens): Total
diameter of test 373–448, diameter of central depression
211–249.
Remarks:P. alaskaensis n. sp. differs from all the other
species of the genus by having the equatorial plate exten-
ded outside the rim to form a porous keel and by having
numerous short, flat, and triangular spines. By this char-
acter, it seems to be close to Carinacyclia Kozur &
Mostler.
Etymology: From its occurrence in Alaska.
Range and occurrence: Latest Carnian (latest Tuvalian
or early Norian), Glenn Shale, east-central Alaska.
Praeorbiculiformella bragini Dumitrica & Tekin n. sp.
(Fig. 12a, b)
Description: Test circular with a very narrow but rela-
tively high rim and 11 to about 14 peripheral spines.
Central cavity very wide, but never complete in our spec-
imens. Rim represented by a circumferential row of wide
pores or by only a circular ridge, in which case the row of
pores is located outside this ridge. Spines short, pyramidal,
usually tricostate. Margin of test in the intervals between
spines straight or slightly convex or concave.
Material: One specimen with complete ring and several
fragments, almost all in the topotype material, sample
BR918, Zulla Formation, Wadi Bani Khalid section,
Oman.
Holotype: Fig. 12a, Muse
´edeGe
´ologie Lausanne, No.
MGL 96843.
384 P. Dumitrica et al.
123
Author's personal copy
Fig. 12 a,bPraeorbiculiformella bragini Dumitrica & Tekin n. sp.,
BR918: aholotype, bparatype, scale =140 lm. c–fPraeorbiculi-
formella gokceoglui Dumitrica & Tekin n. sp.: c,dholotype, 04-ELB-
5, ddetail of the microsphere, scale bar =250 and 15lm;
e,fparatypes: e04-ELB-5, f04-ELB-1, scale bar =250 lm.
g–iPraeorbiculiformella lahmi Dumitrica & Tekin n. sp.: gholotype,
04-ELB-2, scale bar =230 lm; h–iparatypes: h04-ELB-3,
i04-ELB-2, scale bar =230 lm. jPraeorbiculiformella latimargi-
nata Kozur & Mostler, 04-ELB-2, scale bar =230 lm. k,mPrae-
orbiculiformella parvicentrum Dumitrica & Tekin n. sp.: kholotype,
04-ELB-2; l,mparatypes: l04-ELB-2, m04-ELB-1, scale bar for
all =210 lm. n–pPraeorbiculiformella plana Kozur & Mostler,
04-ELB-2, scale bar =210 lm. q,rPraeorbiculiformella polycircus
Dumitrica & Tekin n. sp.: qparatype, 04-ELB-5, rholotype, 04-ELB-
2, scale bar =180 lm. s,tPraeorbiculiformella polyspinosa Kozur
& Mostler, 04-ELB-2, scale bar =220 lm. u,vPraeorbiculiformella
tethyus (De Wever), u04-ELB-1, v04-ELB-2, scale bar =180 lm.
wPraeorbiculiformella sp. A, 04-ELB-1, scale bar =150 lm.
xPraeorbiculiformella sp. B, 04-ELB-2, scale bar =130 lm
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 385
123
Author's personal copy
Dimensions (in lm based on two specimens): Diameter
of test without spines 249–261, length of spines 29–32.
Remarks: This new species is very well distinguished
from all the species of the genus Praeorbiculiformella, so
far described by its very narrow rim and the short spines.
Etymology: The species is named for Dr. Nikita Yu.
Bragin, Moskow, to honour of his contribution to the
knowledge of Mesozoic radiolarians.
Range and occurrence: Early Tuvalian, Spongotortili-
spinus moixi Zone, Zulla Formation, Wadi Bani Khalid
section, Oman.
Praeorbiculiformella gokceoglui Dumitrica & Tekin n. sp.
(Fig. 12c–f)
1999 Orbiculiforma plana (Kozur & Mostler).—Tekin:
119, pl. 21 Fig. 4
Description: Test large, mainly flat with 8–14 long
peripheral spines. Peripheral spines unequally spaced, very
flat, triangular, weakly bladed, with the same pore frames
as those of the rim on the proximal part and needle-like on
the distal one. Interspinal areas usually concave. Rim of the
test wide and very shallow, with irregularly arranged
polygonal (triangular to hexagonal, but mainly hexagonal)
pore frames when thicker, and usually with an imperforate
band on the external border. When thin, rim shows con-
centric rows of pores. Central cavity wide, occupying
approximately 2/3 of the test, with mainly hexagonal pore
frames arranged in four to six concentric circles. Slope
between rim and central cavity very gentle, almost
indistinct.
Material: More than 20 specimens in samples 04-ELB-1
and 04-ELB-5, of which four have been photographed.
Holotype: Fig. 12c, d, Muse
´edeGe
´ologie Lausanne, No.
MGL 96844.
Dimensions (in lm, based on four specimens): Diameter
of test 300–370, diameter of central cavity 185–264, length
of spines 71–142.
Remarks: This new species differs essentially from
Praeorbiculiformella plana Kozur & Mostler by having a
more flattened shell and proximally flat triangular spines.
Etymology: The species is dedicated to Dr. Candan
Gokceoglu (Hacettepe University, Ankara, Turkey) in
honor of his contribution to the study of fuzzy logic
applications in Geology.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Praeorbiculiformella lahmi Dumitrica & Tekin n. sp.
(Fig. 12g–i)
Description: Test large, rather flat with 9–14 long
peripheral spines. Spines unequally spaced, with triangular
base and needle-like distal end. Interspinal areas concave
when spines are closely spaced and straight or very slightly
concave when interspinal space is large. Rim of the test
rather narrow, relatively thick and rather well marked from
the low central cavity. Pores of rim irregularly arranged
and polygonal (triangular to hexagonal, but mainly hex-
agonal). Central cavity wide, occupying approximately 2/3
of test diameter, with mainly hexagonal pore frames
arranged in four to six concentric circles.
Material: Three illustrated specimens from samples
04-ELB-2 and 04-ELB-3.
Holotype: Fig. 12g Muse
´edeGe
´ologie Lausanne, No.
MGL 96845.
Dimensions (in lm, based on three specimens) Diameter
of test 274–334, diameter of central cavity 177–246, length
of spines 70–129.
Remarks: This new species resembles very much Prae-
orbiculiformella gokceoglui n. sp. from which it differs in
having a thicker and rather well defined rim and thinner
spines on which the spongy test of the rim is not prolonged
at all or only on their base.
Etymology: The species is named for Dr. Bernhard
Lahm for their contribution to the knowledge of Triassic
radiolarians.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Praeorbiculiformella latimarginata (Kozur & Mostler
1978)
(Fig. 12j)
1978 Praeorbiculiformella latimarginata Kozur &
Mostler: 165, pl. 4 Fig. 8
1984 Praeorbiculiformella latimarginata Kozur &
Mostler.—Lahm: 93, PL.17FIG.3
Remarks: The species is rather sparse in the Ko
¨seyahya
samples.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Grossreifling, Austria; Ko
¨seyahya
section, Elbistan, SE Turkey.
Praeorbiculiformella parvicentrum Dumitrica & Tekin n. sp.
(Fig. 12k–m)
386 P. Dumitrica et al.
123
Author's personal copy
Description: Large, disc-shaped test with six to eight
peripheral spines, rarely more, a wide and low rim, and a
small central depression. Rim polygonal, flat, rather low,
about half the length of arm in breadth, with pore frames
arranged in three to four concentric circles in the inner half
and irregularly arranged in the outer half. Pore size of rim
increasing distally. Peripheral spines long, conical with
three or more ridges on basal and middle parts, then nee-
dle-like distally. Periphery of rim rounded in transversal
section, straight or slightly concave between spines. Cen-
tral cavity of the test rather well delimited from the rim,
circular, with big polygonal (mainly hexagonal) pore
frames of irregular arrangement. Central part of test small,
raised, consisting of very delicate spongy meshwork.
Material: Four specimens in the samples 04-ELB-1 and
04-ELB-2, of which three have been photographed.
Holotype: Fig. 12k, Muse
´edeGe
´ologie Lausanne, No.
MGL 96846.
Dimensions (in lm, based on three specimens) Outer
diameter of test 291–298, diameter of central cavity
188–198, maximum length of spines 107–110.
Remarks:Praeorbiculiformella parvicentrum n. sp.
resembles very much P. plana Kozur & Mostler in having a
low rim, but differs from it by having a smaller central
cavity, pore frames wider and irregularly disposed in this
cavity, and more robust spines.
Etymology: From the Latin parvus—small and cen-
trum—centre.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Praeorbiculiformella plana Kozur & Mostler 1978
(Fig. 12n–p)
1978 Praeorbiculiformella plana Kozur & Mostler: 164,
pl. 1 Figs. 11, 12, 15; pl. 4 Figs. 1, 2, 4, 9, 10
1984 Praeorbiculiformella plana Kozur & Mostler.—
Lahm: 92, pl. 16 Fig. 11
Description: Test large, circular, biconcave lens-shaped
with a narrow rim and a wide central cavity. Periphery
armed with six to ten, rarely more, commonly eight long to
moderately long, bladed, and pointed spines. Spines nar-
row, equally to unequally spaced, and terminated usually in
a needle-like spike. Pore frames of rim commonly continue
along half the length of spines. Rim of test wide, moder-
ately thickened with irregularly arranged pores. Its pores
are polygonally framed and slightly larger than in central
part. Rim slopes gently toward central cavity, which
occupies about one half to 2/3 of test. Central cavity thin,
double-layered with hexagonally framed pores arranged in
up to ten concentric circles.
Material: Fifteen specimens in sample 04-ELB-2, of
which three photographed.
Dimensions (in lm, based on five specimens): Diameter
of test 274–355, diameter of central cavity 177–258, length
of peripheral spines 50–129.
Remarks: This species differs from P. vulgaris Kozur &
Mostler by having more and longer peripheral spines, rim
with irregular pore frames, gentle slope between rim and
central cavity, and concentrically disposed pores in central
cavity.
Range and occurrence: Early to middle Carnian. Middle
Carnian, Tetraporobrachia haeckeli Zone: Go
¨stling and
Grossreifling, Austria; Ko
¨seyahya section, Elbistan, SE
Turkey; early Carnian, Tritortis kretaensis Zone, to middle
Carnian, Tetraporobrachia haeckeli Zone, Zulla Forma-
tion, Al Jil and Wadi Bani Khalid sections, Oman.
Praeorbiculiformella polycircus Dumitrica & Tekin n. sp.
(Fig. 12q, r)
Description: Test circular to polygonal, usually with
eight to ten or more peripheral spines. Spines rather robust,
pyramidal, usually three-bladed and as long as the breadth
of the rim or a little longer. Rim very narrow but very
thick, with small pores, which are disposed in about two
rather disturbed circular rows and which bear or not very
small nodes at vertices. Boundary with central cavity rather
sharp, practically vertical. Central cavity wide with five to
six circles of pores. Center small and raised, with a dense
spongy meshwork.
Material: Several tens of specimens in samples 04-ELB-
2 and 04-ELB-5, of which two have been photographed.
Holotype: Fig. 12r, Muse
´edeGe
´ologie Lausanne, No.
MGL 96847.
Dimensions (in lm based on two specimens): Diameter
of shell without spines 285–343, diameter of central cavity
200–271, length of spines 57–71.
Remarks: The rim of this species is somehow interme-
diate between that of P. vulgaris Kozur & Mostler and P.
polyspinosa Kozur & Mostler, but the central depression
differs from that of the two species by having the pores
disposed in five to six concentric circles.
Etymology: From the Latin, meaning many circles.
Range and occurrence:MiddleCarnian(Tetraporobra-
chia haeckeli Zone): Ko
¨seyahya section, Elbistan, SE Turkey.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 387
123
Author's personal copy
Praeorbiculiformella polyspinosa Kozur & Mostler 1978
(Figs. 9d, g, 12s, t)
1978 Praeorbiculiformella polyspinosa Kozur &
Mostler: 166, pl. 4 Fig. 11
Description: Test disc-shaped, circular to polygonal,
usually with 8–11 peripheral spines. Spines rather robust,
pyramidal, usually three-bladed and a little longer than the
breadth of rim. Rim very narrow but thick, with small pores
which are irregularly disposed or in two rather disturbed
circular rows, and which bear or not small nodes at verti-
ces. Boundary with central cavity sharp, vertical. Central
cavity thin and wide with pores disorderly arranged. Center
small and raised, with a dense spongy meshwork.
Material: Ten specimens in sample 04-ELB-2, of which
two have been photographed.
Remarks: In the Ko
¨seyahya section, this species is very
rare as compared with the other species of this genus. It can
be easily differentiated from the other species with prac-
tically similar rim by having the pores of the central cavity
disorderly arranged. A similar central cavity has Praeor-
biculiformella vulgaris Kozur & Mostler, but it has the rim
with usually two more or less irregular circular rows of
pores.
Range and occurrence: Middle Carnian (late Julian),
Tetraporobrachia haeckeli Zone: Grossreifling, Austria;
Ko
¨seyahya section, Elbistan, SE Turkey.
Praeorbiculiformella tethyus (De Wever 1979)
(Fig. 12u, v)
1979 Perispongidium (?) tethyus n. sp. De Wever.—De
Wever et al.: 94, pl. 7 Figs. 6–8
1982 Orbiculiforma tethyus (De Wever).—De Wever:
194, pl. 31 Figs. 2–6
1999 Praeorbiculiformella goestlingensis Kozur &
Mostler.—Bragin & Krylov: 554 Fig. 8d
2007 Orbiculiforma goestlingensis (Kozur & Mostler).—
Bragin: 1000, pl. 11 Fig. 8
Description: Test large, circular to slightly polygonal
with a thick, narrow rim and a wide central cavity.
Central cavity thin, usually one-layered with polygonally
framed pores disposed in up to ten concentric circles.
Central shell very small, globular, made up of very thin
meshwork. The four primary beams usually visible, very
thin, aligned to four of the peripheral spines. Peripheral
rim thick and narrow with a vertical or even overhanging
wall at the boundary with central cavity and with pores
disposed in one to two concentric circles, the inner one
well marked, whereas the outer circle is irregular and has
smaller pores. Inner and outer margins of inner circle
with nodes at vertices of pore frames. Usually the nodes
of the inner margin are bigger that those of the outer
margin of the circle. Periphery of rim rounded or
polygonal, and armed with 7–12 spines, most frequently
eight spines. Spines relatively long, thick, pyramidal,
usually three-bladed, rarely terminated in a needle-
shaped shaft.
Remarks: It is strange that De Wever (in De Wever et al.
1979) chose as holotype a specimen in lateral view that
does not show the primary taxonomic character of the
species, which is a face view. It is probably because it was
the only specimen of the two originally presented that was
illustrated in electronic microscopy. Fortunately, the
paratype, illustrated in optic microscopy and showing a
face view, and the subsequent illustration of other two
specimens (De Wever, 1982) from the same sample show
very clearly the taxonomic characters of the species. The
species is very rare in the Ko
¨seyahya samples. The rim of
this species and the number and morphology of spines
somehow resemble those of P. polyspinosa Kozur &
Mostler, from which it differs in having the pores of the
rim disposed much more regularly in one circular row of
pores bordered on both sides (internal and external) by two
rows of nodes. From the poorly illustrated holotype of P.
goestlingensis Kozur & Mostler, the species differs in
having a narrower, nodular rim and pores of rim disposed
in one concentric row.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, to early Norian: late Carnian—early
Norian of Greece; early Norian, Cyprus; middle Carnian,
Ko
¨seyahya section, Elbistan, SE Turkey.
Praeorbiculiformella sp. A
(Fig. 12w)
Remarks: This morphotype, from which we have only
one specimen, resembles partly P. plana Kozur & Mostler,
from which it differs in having fewer rows of pores.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Praeorbiculiformella sp. B
(Fig. 12x)
Remarks: The central depression of this species, with
irregularly disposed pores, resembles that of P. polyspin-
osa, but the peripheral rim is different.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, eastern
Turkey.
388 P. Dumitrica et al.
123
Author's personal copy
2c. Subfamily TRIASSOCRUCELLINAE Dumitrica & Tekin,
n. subfam.
Type genus:Triassocrucella Kozur 1984.
Diagnosis: Veghicycliidae with four spongy arms
developed around and along the four primary beams. Arms
usually terminated in a central spine, which represents
prolongation of the primary beam, and two lateral spines.
Remarks: So far, this subfamily is represented by only
its type genus. By its microsphere and the presence of four
primary beams, it is close to the subfamilies Tetrapauri-
nellinae n. subfam. and Veghicycliinae. It differs from the
former in having a spongy cortical shell around the primary
beams, form the latter in having a shell only around the
primary beams.
Range: Late Triassic.
Genus Triassocrucella Kozur 1984
Type species:Hagiastrum baloghi Kozur & Mostler,
1978.
Remarks: The genus was erected by Kozur (1984)to
differentiate it from Crucella Pessagno. According to him, this
genus differs from Crucella in having bulbous tips with three
to four spines and no patagium. The sections we made in all
species of Triassocrucella support this differentiation and
prove that their microsphere is quite different from that of the
Jurassic and Cretaceous four-armed Hagiastridae as described
and illustrated by P. Dumitrica in De Wever et al. (2001). It
resembles the microsphere of the Tetrapaurinellinae and
Veghicycliinae. Corners of the microsphere are at the origin of
four primary beams. These beams are massive or slightly
tricostate and have numerous branches which are not always
disposed in verticils of three,but turn successively about 120°
as added around the primary beam.
In spite of the similarity between the morphology and
arm structure of the members of this genus and the
Hagiastridae, Triassocrucella is not a forerunner of the
Hagiastridae. These two groups are just a good example of
homeomorphy among radiolaria. The Hagiastridae, as we
shall discuss in another paper arised from Icrioma De
Wever and Paricrioma Tekin.
Range: Carnian to Rhaetian.
Triassocrucella baloghi (Kozur & Mostler 1978)
(Fig. 13a–c)
1978 Hagiastrum baloghi Kozur & Mostler: 144, pl. 2
Figs. 1–5
1984 Crucella baloghi (Kozur & Mostler).—Lahm: 90,
pl. 16 Fig. 7
1990 Crucella baloghi (Kozur & Mostler).—Kolar-
Jurkovsek: 81, pl. 10 Fig. 1
1984 Triassocrucella baloghi (Kozur & Mostler).—
Kozur: 33, pl. 1 Fig. 3
1999 Triassocrucella baloghi (Kozur & Mostler).—
Tekin: 88, pl. 9 Fig. 5
Material: Fourteen specimens in sample 04-ELB-2, of
which three have been photographed.
Remarks: This species shows a rather wide variability,
from small forms with broad distal end armed with three
short spines interconnected by a blade, similar to the
holotype, to larger forms similar to the paratype and the
specimen illustrated by Lahm (1984). The latter forms are
morphologically transitional to T. goestlingensis. Other
forms are morphologically transitional to T. sireli n. sp.
Very common to this species are its small size, the exag-
gerated broadening of the distal ends of arms and, usually,
the appearance of a blade on the arm tip between the
central spine and the two lateral spines. As anomalies, we
mention here one specimen with five arms (Fig. 13b), of
which one is normal, of morphology characteristic of the
species, whereas the other four arms are thinner and
practically symmetrical relative to a perpendicular plan
passing through the normal arm.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone to early Tuvalian, Spongotortilispinus
moixi Zone: Go
¨stling and Grossreifling, Austria; Slovenia;
Bozkir, Konya, and Ko
¨seyahya section, Elbistan, SE Tur-
key; Zulla Formation, Wadi Bani Khalid, Oman.
Triassocrucella goestlingensis (Kozur & Mostler 1978)
(Figs. 9c, 13d–g)
1978 Hagiastrum goestlingense Kozur & Mostler: 144,
pl. 1 Fig. 5, pl. 2 Figs. 6–8
1984 Crucella goestlingensis (Kozur & Mostler).—
Lahm: 90, pl. 16 Fig. 8
1984 Triassocrucella goestlingensis (Kozur &
Mostler).—Kozur: 33, pl. 1 Figs. 1, 4
Material: Tens of specimens.
Remarks: This species is the most frequent species of
Triassocrucella in the Ko
¨seyahya samples. It is very
constant in its morphology. As anomaly we mention
here a rather symmetrical form with five arms (Fig. 13g).
Such anomalies are extremely rare in the genus
Triassocrucella.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling and Großreifling, Austria;
Ko
¨seyahya, eastern Turkey.
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 389
123
Author's personal copy
Fig. 13 a,cTriassocrucella baloghi (Kozur & Mostler), 04-ELB-2,
scale bar =220 lm. d–gTriassocrucella goestlingensis (Kozur &
Mostler): d04-ELB-1, e04-ELB-2, fand g04-ELB-5, scale
bar =230 lm. h–jTriassocrucella kamatai Dumitrica & Tekin n.
sp.: hholotype, 04-ELB-2; i,jparatypes: i04-ELB-2, j04-ELB-5,
scale bar =270 lm. k,lTriassocrucella karnica (Kozur & Mostler):
k04-ELB-1, l04-ELB-2, scale bar =160 lm. m,nTriassocrucella
longispinosa (Kozur & Mostler): m04-ELB-2, n04-ELB-5, scale
bar =190 lm. oTriassocrucella obesa (Kozur & Mostler), 04-ELB-
1, scale bar =170 lm. p–rTriassocrucella sireli Dumitrica & Tekin
n. sp.: pholotype, 04-ELB-5; q,rparatypes, 04-ELB-2 scale bar for
all specimens =200 lm. sTriassocrucella triassica (Kozur &
Mostler), 04-ELB-2, scale bar =290 lm
390 P. Dumitrica et al.
123
Author's personal copy
Triassocrucella kamatai Dumitrica & Tekin n. sp.
(Figs. 9e, 13h–j)
Description: Test with short, very broad, rectangular,
and practically equal arms. Arms elliptical in cross-section,
flat, with parallel or subparallel sides due a slight increase
or decrease in width distally. Cortical test composed of an
irregular network of small, triangular and quadrangular
pore frames with nodes at pore frame vertices. Arm tips
truncate with two or three needle-like and distally directed
short or very short spines; central spine, when present,
slightly longer than the lateral ones or practically absent.
Microsphere small with about 12 wide polygonal pores. It
is at the origin of four primary beams running along the
central part of arms and connected to spongy skeleton of
arms by rather irregular short verticils of three arms con-
tinuously rotated as added, so that they leave no primary
canals.
Material: Eight specimens in samples 04-ELB-2 and
04-ELB-5, of which three have been photographed.
Holotype: Figure 13h, Muse
´edeGe
´ologie Lausanne,
No. MGL 96848.
Dimensions (in lm, based on five specimens): Length of
arms 133–160, width of arms at base 99–117, length of
distal spines 83–100.
Remarks:Triassocrucella kamatai n. sp. very much
resembles T. triassica (Kozur & Mostler), from which it
differs in having arms with parallel to slightly subparallel
sides and truncate tips.
Etymology: The species is dedicated to Dr. Yoshihioto
Kamata (Department of Earth Science, Yamaguchi Uni-
versity, Japan) to honour his contributions to the study of
Triassic radiolarian biostratigraphy.
Range and occurrence:MiddleCarnian,Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE Turkey.
Triassocrucella karnica (Kozur & Mostler 1978)
(Fig. 13k, l)
1978 Hagiastrum karnicum Kozur & Mostler: 145, pl. 1
Fig. 6, pl. 2 Figs. 9, 13
1984 Crucella karnica (Kozur & Mostler).—Lahm: 91,
pl. 16 Fig. 10
Material: Ten specimens.
Remarks: The absence of lateral spines on the tips arms
would suggest that this species is morphologically closer to
Crucella Pessagno than to Triassocrucella Kozur. How-
ever, since no typical microsphere of Crucella-type was
found in the Carnian and Norian species we investigated
and one small lateral spine may by present on some
specimens, we think that this species can undoubtedly be
assigned to Triassocrucella.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Go
¨stling and Grossreifling, Austria;
Ko
¨seyahya section, Elbistan, SE Turkey.
Triassocrucella longispinosa (Kozur & Mostler 1978)
(Fig. 13m, n)
1978 Hagiastrum longispinosum Kozur & Mostler: 145,
pl. 3 Fig. 1
Description: Test small, thick in the central area, sloping
away toward distal ends of arms. Arms short, decreasing in
breadth distally and terminating in a long conical spine. At
its base, this spine may be accompanied by one or two very
short lateral spines. Surface of test with irregularly arran-
ged nodes interconnected by thin bars forming triangular to
pentagonal meshes. Internal structure unknown.
Material: Two specimens, one in 04-ELB 2, and one in
04-ELB 5.
Remarks: Although the morphology of this species is
very close to that of Crucella Pessagno and we do not
know the microsphere due to the poor preservation of the
specimens, the presence of very small lateral spines on
some arms suggests that it belongs to Triassocrucella
Kozur rather than to Crucella. The holotype of the species
shows no lateral spines characteristic of Triassocrucella,
whereas in our material one specimen (Fig. 13m) has very
small lateral spines on two arms and another one (Fig. 13n)
has such spines on all arms. Their presence changes the
shape of arms in face view from conical to rectangular.
Since both specimens have long axial spines characteristic
of the holotype we consider that they represent variation of
the same species.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone; Go
¨stling, Austria; Ko
¨seyahya section,
Elbistan, SE Turkey.
Triassocrucella obesa (Kozur & Mostler 1978)
(Fig. 13o)
1978 Hagiastrum obesum Kozur & Mostler: 146, pl. 3
Fig. 3
1984 Triassocrucella obesa (Kozur & Mostler).—Kozur: 33
Description: Test flat with broad and short arms. Arms
with practically parallel sides and transversally cut ends,
which usually bear two or more very short, radially directed
spines. Central part of test with a very dense spongy mesh-
work. Size of meshwork increases distally along arms, where
Taxonomic study of spongy spumellarian Radiolaria with three and four coplanar spines 391
123
Author's personal copy
it forms four to five radial rows of pores separated by radial
beams and alternately disposed. Interradial areas with a short
patagium of irregularly disposed polygonal meshes.
Material: Two specimens.
Remarks: The species is very rare and is the only species of
this genus that shows the radial structure of the arms on the
surface of shell. The shape of arms resembles that of Trias-
socrucella triassica (Kozur & Mostler), from which it differs
especially in the radial disposition of the pores on the arms.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone: Grossreifling, Austria; Ko
¨seyahya
section, Elbistan, SE Turkey.
Triassocrucella sireli Dumitrica & Tekin n. sp.
(Fig. 13p–r)
Description: Test small with short, deeply forked arms.
Branches of arms conical, cylindrical or irregular, usually
terminated with short spines. Commonly, between the two
arms a short, rarely long spine develops as an external
prolongation of the primary beam. Generally, distal part of
arms is very irregular, the intermediary spine may be
deviated towards one branch or may be missing. Cortical
test composed of polygonal (triangular to pentagonal) or
irregular pore frames of different size and arrangement,
although, partly, a radial disposition may develop.
Material: Fifteen specimens in samples 04-ELB-2 and
04-ELB-5, of which three have been photographed.
Holotype: Fig. 13p, Muse
´edeGe
´ologie Lausanne, No.
MGL 96849.
Dimensions (in lm, based on 10 specimens): Length of
arms 133–166 (average 150), width of arms at the base 55–80
(average 63), length of spines at tips 44–67 (average 57).
Remarks: This species differs of all the other species of
Triassocrucella Kozursofardescribedinhavingbifurcatedor
trifurcated arms. It partly resembles T. baloghi (Kozur &
Mostler), but it is distinguished from this species in having
forked arms, although some intermediary specimens between
them may occur.
Etymology: The species is dedicated to Dr. Ercument Sirel
(Ankara University, Ankara, Turkey) in honor of his contri-
bution to the biostratigraphy of Tertiary Foraminifera.
Range and occurrence: Middle Carnian, Tetraporobra-
chia haeckeli Zone, Ko
¨seyahya section, Elbistan, SE
Turkey.
Triassocrucella triassica (Kozur & Mostler 1978)
(Fig. 13s)
1978 Hagiastrum triassicum Kozur & Mostler: 146, pl. 1
Fig. 4; pl. 2 Fig. 11, ?12; pl. 3 Fig. 2
1984 Crucella triassica (Kozur & Mostler).—Lahm: 91,
pl. 16 Fig. 9
1984 Triassocrucella triassica (Kozur & Mostler).—
Kozur: 33
1990 Hagiastrum triassicum Kozur & Mostler.—Budai
& Dosztaly, pl. 3 Fig. 1
1993 ?Triassocrucella sp. aff. T. triassicum (Kozur &
Mostler).—Carter: 85, pl. 11 Fig. 16
1999 Triassocrucella triassica (Kozur & Mostler).—
Tekin: 88, pl. 9 Fig. 6
1999 Triassocrucella triassica (Kozur & Mostler).—
Bragin & Krylov: 554 Fig. 8J
2007 Triassocrucella triassica (Kozur & Mostler).—
Bragin: 1003, pl. 12 Fig. 2
Remarks: The species very much resembles T. goes-
tlingensis (Kozur & Mostler) from which it only differs in
having broader arms. The best-illustrated specimen to
which we should refer for the determination of this species
is the topotype specimen illustrated by Lahm (1984)
because the holotype, illustrated under binocular, shows no
detail of the surface, and the paratypes are either badly
preserved or anomalous. In the Ko
¨seyahya fauna, the
species is very rare and represented by specimens resem-
bling more or less the holotype. The early Norian specimen
illustrated by Bragin & Krylov (1999) and Bragin (2007)is
also rather close to the holotype, whereas the Rhaetian
specimen illustrated by Carter (1993) has slender arms and
is morphologically intermediate between T. triassica and
T. goestlingensis. The species is also compared with
T. kamatai n. sp. under the latter species.
Material: Five specimens, of which one, from sample
04-ELB-2, was photographed.
Range and occurrence: Middle Carnian—Rhaetian;
Go
¨stling and Großreifling, Austria; Kemer area and
Ko
¨seyahya section, Turkey; Mamonia Complex, Cyprus;
Sandilands Formation, Queen Charlotte Islands.
Conclusions
The taxonomy at generic and suprageneric levels, primarily
based on the types of microspheres as the first and most
conservative skeletal element, allowed new taxonomic
arrangements of genera and suprageneric units and sug-
gested new and unexpected phylogenetic relationships
among these radiolarians and between them and younger
radiolarians. Among other results, this study suggests that:
392 P. Dumitrica et al.
123
Author's personal copy
a) during the middle Triassic there was a trend towards
the increase of the complexity of skeletal projections
from cylindrical or conical spines to three-bladed
spines and finally to spongy arms;
b) the origin of the family Tritrabidae is in the three-
spined spongy middle Triassic radiolarians of the
subfamily Intermediellinae;
c) there is a close relationship between the four-spined
lens-shaped spongy radiolarians assigned to the sub-
family Tetrapaurinellinae and the flat circular radiola-
rians of Veghicyclia and Praeorbiculiforma type, and
finally the four-armed radiolarians of the genus
Triassocrucella;
d) the origin of latest Triassic and post-Triassic Hagia-
stridae is not in the genus Triassocrucella as thought
before, but in the genera Icrioma and Paricrioma, as
we will show in a forthcoming paper.
One of the most important conclusions of this study is
that the knowledge of the microsphere and of the first shells
of Spumellaria should represent the basis of the taxonomy
at generic and suprageneric levels of these radiolarians.
The authors discuss and reject the biostratigraphic
Elbistanium gracile Zone described by Kozur et al. 2009
on the basis of the radiolarian assemblage from the
Ko
¨seyahya section.
Acknowledgments All the photos used in this study have been taken
with the CamScan electronic microscope of the Hochschule der Ku
¨nste
Bern (High School of Art of Bern), Switzerland. P. Dumitrica is very
much obliged to Dr. Nadim C. Scherrer, for his kindness and permission
to use this microscope any time he needed it. The senior coauthor thanks
also his wife, Ruth Dumitrica, for the translation of the abstract into
German. The authors also very much thank the two reviewers: Spela
Gorican, Ljubljana, and Luis O’Dogherty, Cadiz, for their critical
reading of the manuscript and very constructive suggestions.
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