GEOLOGICA CARPATHICA, 53, 6, BRATISLAVA, DECEMBER 2002
369 — 379
CHITINOIDELLA
TREJO, 1975
IN MIDDLE TITHONIAN CARBONATE PELAGIC SEQUENCES
OF THE WEST CARPATHIAN TETHYAN AREA
DANIELA REHÁKOVÁ
Geological Institute of the Slovak Academy of Sciences, Dúbravská cesta 9, 842 26 Bratislava, Slovak Republic
Department of Geology and Palaeontology, Faculty of Science, Comenius University, Mlynská dolina G 1,
842 12 Bratislava, Slovak Republic; rehakova@fns.uniba.sk
(Manuscript received March 19, 2002; accepeted in revised form June 18, 2002)
Abstract: Detailed microfacies analysis of Tithonian pelagic carbonate sequences from the Central West Carpathian and
Pieniny Klippen Belt areas allows us to recognize several differences in chitinoidellid morphology on the basis of which
several chitinoidellid genera – Chitinoidella Doben, 1963; Borziella Pop, 1997; Carpathella Pop, 1998a; Longicollaria
Pop, 1997; Daciella Pop, 1998a; and Dobeniella Pop, 1997 could be recognized among the Middle Tithonian chitinoidellids
of the Dobeni and Boneti Subzones. The new Popiella n.gen. was introduced, and in the framework of it, the new
Popiella oblongata n.sp. was described. On the basis of the chitinoidellid vertical distribution, they were included into
the Dobeni and the Boneti Subzones of the standard Chitinoidella Zone where they can be utilized for High Resolution
Stratigraphy (HIRES) of the Middle Tithonian carbonate pelagic sequences. From the point of view of chitinoidellid
phylogeny, this paper regards Longicollaria dobeni as the ancestor of Dobeniella and Crassicollaria species; Dobeniella
tithonica could have been the ancestor of Remaniella ferasini. The newly described Popiella oblongata could have been
in phylogenetic lineage with Calpionellopsis simplex and Calpionellopsis oblonga.
Key words: Tithonian, Western Carpathians, Tethys, evolution, phylogeny, taxonomy, biozonation, bioevents,
chitinoidellids.
Introduction
Chitinoidellids represent a very small group of ancient plank-
tonic ciliates with microgranular, axially symmetrical lorica.
The genus Chitinoidella was established by Doben (1962). It
comprised all ciliate forms showing a dark wall in transmitted
light. For this reason, Doben supposed that chitinoidellid wall
is built of chitinous matter. Borza (1966, 1969) and Remane
(1969) stated that chitinoidellids have microgranular calcitic
loricas. Later, Reháková & Michalík (1992, 1993) confirmed
this hypothesis in their studies of the chitinoidella wall ultra-
structure by SEM method. Their observations showed that the
wall of chitinoidellid lorica is composed of three zones. The
two outer zones are formed by scalenohedral calcite crystal-
lites perpendicular to the lorica wall, while the central dark
zone consists of small rhombohedral transversely oriented
calcite crystallites and also of remains of organic substance.
Chitinoidellids were formerly assigned to the single genus
Chitinoidella Pop (1997, 1998a) made their taxonomic revi-
sion and regrouped them into several new genera. The author
of this contribution has tried to revise previous knowledge on
these microplanktonic organisms from the West Carpathian
area and to correlate these results with those stated by Pop
(1997, 1998a). She has tried to bring further lights to the tax-
onomy and phylogeny of the chitinoidellids.
Geological and lithological background
Jurassic
and
Lower
Cretaceous
formations
form
a considerable part of the Mesozoic sequences in both, the
Outer and Central Western Carpathians (Fig. 1). Upper Juras-
sic—Lower Cretaceous carbonate microfacies and environ-
mental models for the Western Carpathians and adjacent pa-
leogeographic units were created in detail by Vašíček et al.
(1994) and Reháková (1995a). Chitinoidellids are common el-
ements of “Ammonitico Rosso” – type condensed deposits
accumulated on the thresholds in a pelagic environments. The
Czorsztyn Limestone Formation (locally Niedzica or Tegern-
see Limestone Formations), the typical West Carpathian rep-
resentatives of “Ammonitico Rosso” facies consists of Kim-
meridgian and Tithonian reddish nodular or pseudonodular
limestone. They are of biomicrite (packstone) and biomi-
crosparite (wackestone) character. Intraclasts occur frequent-
ly, some beds pass to limestone microbreccia. A rich associa-
tion of microfossils allows us to distinguish several
calpionellid and dinoflagellate zones in the framework of this
limestone sequence: the Kimmeridgian Moluccana and Borzai
Zones, Early Tithonian Pulla, Tithonica and Malmica Zones,
Middle Tithonian Semiradiata, Tenuis and Chitinoidella
Zones and Late Tithonian Fortis, Proxima, Praetintinnopsella
and Crassicollaria Zones (Michalík et al. 1990; Reháková
370 REHÁKOVÁ
2000a,b). In both the Outer and Central Carpathians, con-
densed nodular limestone facies continued on “pelagic eleva-
tions” (Czorsztyn Limestone Formation, Horné Sŕnie Forma-
tion), or shallow marine ridges (Olivetská, Lysa, Spiš and
Raptawicka Turnia Formations) up to the Late Berriasian (Re-
háková 1995a).
Systematics – chitinoidellid genera and species
from the Western Carpathians
On the basis of chitinoidellid morphological features (lorica
and collar construction) it was possible to recognize following
chitinoidellid genera and species.
Family Chitinoidellidae Trejo, 1975
Genus Chitinoidella Doben, 1963
Chitinoidella boneti Doben, 1963
Fig. 2.1—4
1963 Chitinoidella boneti n.sp. – Doben, pl. 6, figs. 1—5
1965 Tintinnopsella carpathica (Murgeanu et Filipescu) – Furrazo-
la-Bermudez, pl. 4, fig. 2
1965 Chitinoidella boneti Doben – Borza, pl. I, figs. 1—6; pl. II,
figs. 1—4
1975 Tintinnopsella carpathica (Murgeanu et Filipescu) – Trejo, pl.
XII, fig. 34
1993 Chitinoidella boneti Doben – Lakova, pl. I, figs. 7—8
1995 Chitinoidella boneti Doben – Reháková, pl. I, figs. 6—7
1995 Chitinoidella boneti Doben – Oloriz et al. pl. I, fig. 3
1998 Chitinoidella boneti Doben – Pop, pl. I, figs. 34—37, 39
D i a g n o s i s : Microgranular calcitic, bell-shaped lorica,
commonly with a slightly preoral constriction and a length/
width ratio smaller than 1.5. Aboral pole of lorica ends by
a short caudal appendage, its large oral opening bears a collar
which outwardly deflected the lorica. Common dimensions
are 55—83 µm in length and 40—50 µm in width. It resembles
that of Tintinnopsella carpathica (Murgeanu et Filipescu) or
Praetintinnopsella andrusovi Borza.
O c c u r r e n c e : Boneti Subzone. Czorsztyn and Tegernsee
Limestone Formations of the Western Carpathians; South
Carpathians, Western Balkanides, NW Anatolia, in the Span-
ish Betic Cordillera and in the Venetian Alps, Eastern Alps,
Cuba, Mexico.
Chitinoidella elongata Pop, 1997
Fig. 2.5—8
1969 Chitinoidella boneti Doben – Borza, pl. LXVIII, fig. 8
1997 Chitinoidella elongata n.sp. – Pop, Fig. 2, photos 3, 4
D i a g n o s i s : Cylindrical or cylindroid lorica with
a conical aboral pole ended by caudal appendage and an out-
Fig. 1. Location of the section studied in the Upper Jurassic and Lower Cretaceous sequence of the Western Carpathians.
CHITINOIDELLA TREJO, 1975 IN TITHONIAN PELAGIC SEQUENCES 371
Fig. 2. 1—4. Chitinoidella boneti Doben. 1 – Manínska úžina, thin section 5472; 2 – Rudník, thin section 5029; 3 – Manínska úžina, thin
section 5472; 4 – Butkov Gallery 13, thin section 8421. 5—8. Chitinoidella elongata Pop. 5 – Manínska úžina, thin section 5472; 6 – Pod-
biel, thin section 4942; 7 – Podbiel, thin section 4941; 8 – Manínska úžina, thin section 5472. 9—12. Borziella slovenica (Borza). 9 – Pod-
biel, thin section 4930; 10 – Podbiel, thin section 4928; 11 – Podbiel, thin section 4930; 12 – Podbiel, thin section 4928. 13—16. Car-
pathella rumanica Pop. 13 – Podbiel, thin section 4930; 14 – Podbiel, thin section 4932; 15 – Podbiel, thin section 4930; 16 – Podbiel,
thin section 4929. 17—20. Daciella danubica Pop. 17 – Podbiel, thin section 4928; 18—19 – Brodno, thin section 5048; 20 – Brodno, thin
section 5047. All figures are related to the bar in Fig. 20 = 100 µm.
372 REHÁKOVÁ
wardly deflected collar. The lorica length is 84—105 µm, its
width is 44—55 µm. Its shape resembles that of Tintinnopsella
longa (Colom).
O c c u r r e n c e : Boneti Subzone. Czorsztyn and Tegernsee
Limestone Formations of the Western Carpathians; Southern
Carpathians.
Genus Borziella Pop, 1997
Borziella slovenica (Borza, 1969)
Fig. 2.9—12
1969 Chitinoidella slovenica n.sp. – Borza, pl. LXVI, figs. 8—9
1993 Chitinoidella slovenica Borza – Lakova, pl. I, fig. 2
1995 Chitinoidella slovenica Borza – Reháková, pl. I, fig. 3
1997 Borziella slovenica (Borza) – Pop, Fig. 2, photos 14—15
1998 Borziella slovenica (Borza) – Pop, pl. I, figs. 16, 17
D i a g n o s i s : Ovoidal to spaeroidal lorica with rounded ab-
oral pole. The lorica length is 40—48 µm, its width is 28—
32 µm. Its preoral part bears a small constriction following by
a relatively short outwardly deflected collar similar to Tintin-
nopsella remanei Borza or Lorenziella hungarica Knauer
type.
O c c u r r e n c e : Dobeni Subzone. Czorsztyn and Tegern-
see Limestone Formations of the Western Carpathians; South-
ern Carpathians, Western Balkanides, NW Anatolia, Eastern
Alps.
Genus Carpathella Pop, 1998
Carpathella rumanica Pop, 1998
Fig. 2.13—16
1969 Chitinoidella sp. – Borza, pl. LXIX, fig. 3
1998 Chitinoidella rumanica n.sp. – Pop, Figure 2, photos 1—5
D i a g n o s i s : Species with globular to ovoidal lorica. Its
bowl is rounded, aboral pole and a slightly constricted preoral
part forming a characteristic “shoulder”. Oral opening contin-
ues by a short cylindrical collar the diameter of which is com-
monly smaller than the maximum width of the lorica. The lor-
ica length is 38—42 µm, its width is 34—36 µm. This species
strongly resembles Calpionella alpina Lorenz.
O c c u r r e n c e : Dobeni Subzone. Czorsztyn Limestone
Formation of the Western Carpathians; Southern Carpathians.
Genus Daciella Pop, 1998
Daciella danubica Pop, 1998
Fig. 2.17—20
1969 Chitinoidella sp. – Borza, pl. LXIX, fig. 4
1998 Daciella danubica n.sp. – Pop, Figure 2, photos 14—18
D i a g n o s i s : Conical and more or less elongated lorica,
commonly with a sharp aboral pole ended by a caudal ap-
pendage and a large opening, preceded by a characteristic
swelling, sometimes hardly perceptible; the oral end of the
lorica shows occasionally a very slight constriction, being
gradually followed by a short cylindrical collar. The lorica
length is 44—58 µm, its width is 34—36 µm. This species
strongly resembles Crassicollaria parvula Remane.
O c c u r r e n c e : Dobeni Subzone. Czorsztyn and Tegern-
see Limestone Formations of the Western Carpathians; South-
ern Carpathians.
Daciella svinitensis Pop, 1998
Fig. 3.1—3
1998 Daciella svinitensis n.sp. – Pop, Fig. 2, photos 19—20
D i a g n o s i s : Lorica elongated or cylindrical its aboral part
ended by a short caudal appendage. The lorica length is 52—
67 µm, its width is 42—45 µm. The large oral opening is sur-
rounded by a short cylindrical collar; the preoral part of the
lorica displays a more or less pronounced swelling resembling
the lorica of Crassicollaria massutiniana (Colom, 1948).
O c c u r r e n c e : Upper part of Dobeni Subzone. Czorsztyn
Limestone Formation of the Western Carpathians; Southern
Carpathians.
Genus Dobeniella Pop, 1997
Dobeniella cubensis (Furrazola-Bermudez, 1965)
Fig. 3.4—6
1965 Tintinnopsella cubensis n.sp. – Furrazola-Bermudez, pl. 1,
figs. 1a—c; pl. 2, figs. 1—5; pl. 5, fig. 1
1966 Chitinoidella cubensis (Furrazola-Bermudez) – Borza, pl. X,
Fig. 10
1995 Chitinoidella cubensis (Furrazola-Bermudez) – Reháková, pl.
I, fig. 10
1998 Dobeniella cubensis (Furrazola-Bermudez) – Pop, pl. I, figs.
27—29
D i a g n o s i s : Bell-shaped, elongated lorica, the oral part of
which ends with a composite collar. The outer ring of collar
is relatively short outwardly deflected, its margin is denticu-
late, the inner ring piece is two times stronger, it is rounded
and more or less closes the lorica oral zone. Aboral pole
pointed ended by a caudal appendage. The lorica length is 55—
72 µm, its width is 45—52 µm.
O c c u r r e n c e : Boneti Subzone. Czorsztyn Limestone
Formation of the Western Carpathians; Cuba, Mexico, South-
ern Carpathians.
Dobeniella bermudezi (Furrazola-Bermudez, 1965)
Fig. 3.7—9
1965 Tintinnopsella bermudezi n.sp. – Furrazola-Bermudez, pl. 1,
figs. 2a—c; pl. 2, figs. 6, 8; pl. 3, fig. 1; pl. 5, fig. 2
1966 Chitinoidella bermudezi (Furrazola-Bermudez) – Borza, pl. X,
Fig. 11
1995 Chitinoidella bermudezi (Furrazola-Bermudez) – Reháková,
pl. I, fig. 9
1998 Dobeniella bermudezi (Furrazola-Bermudez) – Pop, pl. I, figs.
22—24
D i a g n o s i s : The bell-shaped, elongated to subcylindrical
lorica with an aboral pole ended by short pointed caudal ap-
pendage. Uppermost part of lorica body bears the shoulder
following the coarse swelled collar. Sharp distinct steems are
visible on the outer surface of the collar surrounding the mid-
dle part of the swell. The lorica length is 57—85 µm, its width
is 42—57 µm.
CHITINOIDELLA TREJO, 1975 IN TITHONIAN PELAGIC SEQUENCES 373
Fig. 3. 1—3. Daciella svinitensis Pop – all species derived from locality Podbiel, thin section 4913. 4—6. Dobeniella cubensis (Furrazola-
Bermudez). 4 – Brodno, thin section 5061; 5 – Rudník, thin section 5017; 6 – Brodno, thin section 5060. 7—9. Dobeniella bermudezi
(Furrazola-Bermudez). 7 – Manínska úžina, thin section 5471; 8 – Manínska úžina, thin section 5472; 9 – Manínska úžina, thin sec-
tion 5472. 10—12. Dobeniella tithonica (Borza). 10 – Rudník, thin section 2678; 11 – Brodno, thin section 5053; 12 – Brodno, thin
section 5052. All figures are related to the bar in Fig. 12 = 100 µm.
374 REHÁKOVÁ
O c c u r r e n c e : Boneti Subzone. Czorsztyn and Tegernsee
Limestone Formations of the Western Carpathians; Cuba,
Mexico, Southern Carpathians.
Dobeniella tithonica (Borza, 1969)
Fig. 3.10—12
1969 Chitinoidella tithonica n.sp. – Borza, pl. LXVII, figs. 1—2
1993 Chitinoidella tithonica Borza – Lakova, pl. I, fig. 3
1995 Chitinoidella tithonica Borza – Reháková, pl. I, fig. 5
1998 Dobeniella tithonica (Borza) – Pop, pl. I, figs. 25—26
D i a g n o s i s : Shortly elipsoid or bell-shaped lorica bowl.
Its aboral part is pointed or ended by a caudal appendage. An
acute constriction is visible in the preoral part of lorica. Oral
zone of lorica is large. It bears the compose collar consisting
of two rings. The outer, little bit longer ring is outwardly de-
flected in its distal part; the inner one is short and shows lens-
like profile in the cross-section. The lorica length is 42—54 µm,
its width is 34—38 µm.
O c c u r r e n c e : Uppermost part of Dobeni Subzone.
Czorsztyn Limestone Formation of the Western Carpathians;
Southern Carpathians, Western Balkanides.
Dobeniella colomi (Borza, 1966)
Fig. 4.1—3
1966 Chitinoidella colomi n.sp. – Borza, pl. X, figs. 4—7
1995 Chitinoidella colomi Borza – Reháková, pl. I, fig. 4
1997 Dobeniella tithonica (Borza) – Pop, Fig. 2, photo 9
D i a g n o s i s : Bell-shaped lorica ended by a caudal append-
age. Oral belt of lorica is wide, it is followed by distinctly de-
veloped outwardly deflected collar. The wall of the collar is
progressively thickened towards its distal end. The length of
lorica attains 43—60 µm, its width is 28—35 µm; the thickness
of the lorica wall is 2—5 µm.
Species occurs in the uppermost part of the Dobeni Sub-
zone of the Middle Tithonian Chitinoidella Zone.
O c c u r r e n c e : Dobeni Subzone. Czorsztyn Limestone
Formation of the Western Carpathians; Southern Car-
pathians.
Genus Longicolaria Pop, 1997
Longicolaria dobeni (Borza, 1966)
Fig. 4.4—6
1966 Chitinoidella dobeni n.sp. – Borza, pl. IX, figs. 1, 2
1993 Chitinoidella dobeni Borza – Lakova, pl. I, figs. 5, 6
1995 Chitinoidella dobeni Borza – Reháková, pl. I, fig. 2
1997 Longicolaria dobeni (Borza) – Pop, Fig. 2, photos 12—13
1998 Longicolaria dobeni (Borza) – Pop, pl. I, figs. 1—4
D i a g n o s i s : Cup-shaped or ovoid lorica with conical abo-
ral pole ended by a caudal appendage. The preoral segment of
lorica bears a slight constriction following to an elongated cy-
lindrical or subcylindrical collar the wall of which is progres-
sivelly thickened towards the distal end. Its length is compara-
ble with that of the lorica. The lorica length is 48—65 µm, its
width is 34—42 µm.
O c c u r r e n c e : Boneti Subzone. Czorsztyn and Tegernsee
Limestone Formations of the Western Carpathians; Southern
Carpathians, Western Balkanides.
Longicolaria insueta (Řehánek, 1986)
Fig. 4.7—9
1986 Chitinoidella insueta n.sp. – Řehánek, pl. I, figs. 1—4
1993 Chitinoidella insueta Řehánek – Lakova, pl. I, figs. 1
1995 Chitinoidella insueta Řehánek – Reháková, pl. I, 8
1997 Cylindrella insueta (Řehánek) – Pop, Fig. 2, photos 16
1998 Aninella insueta (Řehánek) – Pop, pl. I, figs. 21
D i a g n o s i s : Lorica bowl is elongated vase-like with
a rounded aboral pole, the caudal appendage was not ob-
served. The total lorica length is from 85 to 137 µm, its width
varies between 58—68 µm. The preoral segment of lorica is
without constriction, followed by elongated cylindrical or tab-
ular collar with its wall progressively thickened towards the
distal end. Compared with the lorica bowl, the length of the
collar is two times longer. Lorica shape resembles the colo-
miellid forms.
R e m a r k s : Pop (1997) introduced genus Cylindrella for
ovoidal chitinoidellids showing a rounded aboral pole and
long cylindrical collar. Author described Chitinoidella insueta
Řehánek as its type species. Later Pop (1998b) mentioned,
that the generic name Cylindrella is a homonym (Cylindrella
Sharp, fide Slipinski, 1992) and the author replaced it with the
new generic name Aninella. According to this investigation,
the lorica and collar structure of this species have similar char-
acter to forms belonging to Longicollaria.
O c c u r r e n c e : Boneti Subzone. Czorsztyn Limestone
Formation of the Western Carpathians; Southern Carpathians,
Western Balkanides.
Genus Popiella n.gen.
Lorica bowl is elongated with a rounded aboral pole; the
caudal appendage was not observed. The total lorica length
varies between 65 and 89 µm, its width between 31—38 µm.
The preoral segment of lorica widely opened, sharply ended;
collar was not observed. Lorica shape resembles the forms be-
longing to Calpionellopsis.
Genotype: Popiella oblongata n.gen. n.sp.
Popiella oblongata n.sp.
Fig. 4.10—12
1998 Daciella svinitensis – Pop, Figure 2, Photo 19.
H o l o t y p u s : Specimen on Fig. 4.10, deposited in the
Geological Institute of Slovak Academy of Sciences. Locality
Podbiel, thin section No. 4918.
P a r a t y p e s : Fig. 4.11—12. Locality Podbiel, thin section
4920, 4919.
D e r i v a t i o n o m i n i s : The generic name is devoted to
memory of Grigore Pop, famous micropaleontologist, special-
ist in calpionellids. The name of the species is derived from
the lorica shape.
CHITINOIDELLA TREJO, 1975 IN TITHONIAN PELAGIC SEQUENCES 375
Fig. 4. 1—3. Dobeniella colomi (Borza). 1 – Podbiel, thin section 4929. 2—3. Podbiel, thin section 4930. 4—6. Longicollaria dobeni (Borza).
4 – Podbiel, thin section 4932; 5 – Butkov, Gallery 13, thin section 8423; 6 – Manínska úžina, thin section 5466. 7—9. Longicollaria in-
sueta (Řehánek). 7 – Podbiel, thin section 4941; 8 – Rudník, thin section 5016; 9 – Brodno, thin section 5061. 10. Popiella oblongata
n.sp. (holotype). Podbiel, thin section 4918. 11—12. Popiella oblongata n.sp. (paratypes). 11 – Podbiel, thin section 4920; 12 – Podbiel,
thin section 4919. All figures are related to the bar in Fig. 12 = 100 µm.
376 REHÁKOVÁ
S t r a t u m t y p i c u m : Middle Tithonian, biomicritic nod-
ular, or pseudonodular limestone of the Chitinoidella Zone.
L o c u s t y p i c u s : Klippe Podbiel, Middle Tithonian part
of the Czorsztyn Limestone Formation, Pieniny Klippen Belt.
M a t e r i a l : 5 specimens in thin sections No. 4918, 4919
and No. 4920 (Klippe Podbiel), 2 specimens in thin section
No. 29 (Czorsztyn Limestone Formation of the the Hlboč
section,Vysoká Unit of the Krížna Nappe, Central Western
Carpathians).
D i a g n o s i s : Lorica bowl is elongated with a rounded ab-
oral pole; the caudal appendage was not observed. The total
lorica length (H) varies between 65 and 89 µm, its width (W)
between 31—38 µm. The width of the aperture (A) commonly
attains the lorica maximum width, but sporadically it can be
slightly closer. The wall thickness (Tw) varies between 5—6 µm.
The preoral segment of lorica is widely opened and sharply
ended; collar was not observed. Lorica shape resembles the
forms belonging to Calpionelopsis.
P a r a m e t e r s o f t h e h o l o t y p e : H = 79 µm, W =
35 µm, A = 29 µm, Tw = 6 µm.
A s s o c i a t i o n s : Chitinoidellids of the Boneti Subzone of
the Chitinoidella Zone – Longicolaria insueta (Řehánek),
Chitinoidella boneti Doben 1963, Chitinoidella elongata Pop
in association with abundant Saccocoma Agassiz as well as
Globochaete alpina Lombard, Colomisphaera carpathica
Borza, Schizosphaerella minutissima (Colom), crinoids, bi-
valves, ostracods, aptychi fragments and juvenile ammonites.
O c c u r r e n c e : Boneti Subzone. Czorsztyn and Tegernsee
Limestone Formations of the Western Carpathians; Southern
Carpathians.
Chitinoidellids – stratigraphic distribution and
biozonation
Chitinoidellids attained a dominant position in the plankton
during the Middle Tithonian. Chitinoidella Zone was intro-
duced by Enay & Geyssant (1975) in Spanish Betic Cordillera
and defined by Grandesso (1977) in the Venetian Alps. The
Dobeni and Boneti Subzones (Borza 1984) were characterized
by the total range of the index species. However, Reháková
(1995b) pointed out that the Chitinoidella Zone and both its
subzones are close to interval zones, their upper boundaries
being defined by the first index occurrences of the overlying
younger biostratigraphic unit. The Chitinoidella Zone was
distinguished in the Eastern Alps (Reháková et al. 1996), in
Western Carpathians (Borza 1984; Reháková 1995b), South-
ern Carpathians (Pop 1986, 1989, 1997, 1998a,b), Western
Balkanides (Bakalova 1977; Lakova 1993) and in NW Anato-
lia (Altiner & Özkan 1991). This zone has not been recog-
nized in the Hellenides (Skourtsis-Coroneou & Solakius
1999), Western Mediterranean and westernmost parts of the
Tethyan region (Adatte et al. 1992; Remane et al. 1986). An-
other subdivision of this zone was given by Řehánek (1990).
Consequently, Chitinoidella Zone is defined by the first oc-
currence (FO) of microgranular chitinoidellids. FO of Longi-
collaria dobeni (Borza) represents the beginning of the Dobe-
ni Subzone (the zonal species is accompanied by Longicollar-
ia insueta (Řehánek), Borziella slovenica (Borza), Carpathel-
la rumanica Pop, Daciella danubica Pop, Daciella svinitensis
Pop higher also Dobeniella tithonica (Borza)). Daciella ba-
natica Pop, Daciella almajica Pop and Almajella cristobalen-
sis (Furrazola-Bermudez) were not recognized in the Western
Carpathians section studied.
The FO of Chitinoidella boneti Doben is typical of the
Boneti Subzone. Besides the zonal species, chitinoidellid as-
sociation consists of Chitinoidella elongata Pop, Longicolla-
ria insueta (Řehánek), Dobeniella bermudezi (Furrazola-Ber-
mudez), Dobeniella cubensis (Furrazola-Bermudez), Popiella
oblongata n.sp. as well as some forms persisting from the pre-
ceding interval the Dobeni Subzone.
Chitinoidellid phylogeny
Paleozoic and Triassic microfossils attributed to ancestors
of the calpionellids have been described by Colom (1988);
Visscher (1970, 1971) and Eshet (1990). However, true calpi-
onellid ancestors still remain uncertain. The start of the phylo-
genetic history of calpionellids was connected with rapid radi-
ation of microgranular forms at the beginning of the Middle
Tithonian (Reháková & Michalík 1997; Pop 1997, 1998a).
The evolution of the small chitinoidellids of the Dobeni Sub-
zone followed by the radiation of more advanced and diversi-
fied forms in the Boneti Subzone was finished by chitinoidel-
lid exctinction and their substitution by hyaline forms of the
Praetintinnopsella Zone having an inner hyaline and an outer
microgranular wall layer. The appearence of hyaline calpi-
onellids coincides with the rapid evolution of nannoplankton
associations (Reháková 2002).
Reháková & Michalík (1997) pointed out, that calcite com-
position of calpionellid loricas changed several times during
their evolution. At the beginning of the Middle Aptian new
forms of microgranular praecolomiellids appeared. The verti-
cal span of these specified forms is short and microgranular
loricas are less frequent. On the other hand, they are twice or
several times larger than those of Middle Tithonian chiti-
noidellids. It seems, that the nomismogenesis of planktonic
foraminifers lowered the selectional stress among praecolo-
miellids and such competitive environment led to a growth
expansion of their loricas. During the Early Albian, the prae-
colomiellids were replaced by a new group of hyaline colo-
miellids. The change of lorica composition was synchroneous
with the early Albian peak in nannoconid abundance, as dur-
ing previously described Late Tithonian change of a chiti-
noidellid microgranular structure. On the basis of the first
study of lorica structure in cathodoluminiscence microscope
(Reháková 2000b) considered that during the periods of nan-
noconid blooming tintinnids started to agglutinate their loricas.
Tightly packed coccoliths could have been diagenetically re-
placed by blocky calcite crystals building fossil hyaline spe-
cies. The revival of microgranular calpionellids also allows us
CHITINOIDELLA TREJO, 1975 IN TITHONIAN PELAGIC SEQUENCES 377
Fig. 5. Stratigraphic distribution and possible lineages in the evolution of chitinoidellids. Geochronological data according to Gradstein &
Ogg (1996), actual calpionellid zonation for the Western Carpathians according to Reháková & Michalík (1997).
to speculate about similar climatic and paleoceanographic
conditions during both time intervals.
Pop (1998a) discussed in detail the phylogeny of chiti-
noidellids. He found out, that practically all newly defined
species (Pop 1997) are morphologically very comparable with
some calpionellids of the Crassicollaria and Calpionella
Zones. Among the chitinoidellid associations Pop (1998b)
identified several logical parallel phylogenetic lineages prov-
ing the phylogenetic relationship between chitinoidellids and
calpionellids: Borziella slovenica
→
Tintinnopsella remanei
or Lorenziella hungarica; Carpathella rumanica
→
Calpi-
onella alpina; Daciella danubica
→
Crassicollaria parvula;
Daciella svinitensis
→
Crassicollaria massutiniana; Chiti-
noidella boneti
→
Praetintinnopsella andrusovi
→
Tintin-
nopsella carpathica. All these stocks confirm the phylogenet-
ic concept introduced by Reháková & Michalík (1997).
In contrast to Pop results, this paper (Fig. 5) considers that
Longicollaria dobeni gave rise to stock represented by two
lineages passing through larger Dobeniella species and termi-
nated by Crassicollaria loricas with more-or-less pronounced
swellings below the base of their collars: Dobeniella colomi
→
Crassicollaria intermedia.
378 REHÁKOVÁ
The species Dobeniella tithonica could have been the an-
cestor of the calpionellids with collars composed of one or
several plicae: Dobeniella tithonica
→
Remaniella ferasini.
The newly described Popiella oblongata could have been the
ancestor of the Calpionellopsis type forms: Popiella oblonga-
ta
→
Calpionellopsis simplex
→
Calpionellopsis oblonga.
Conclusions
Detailed investigations of chitinoidellid morphological fea-
tures (lorica and collar construction) allowed us to distinguish
several chitinoidellid genera and species among the Middle
Tithonian chitinoidellids of the Chitinoidella Zone (the Dobe-
ni and the Boneti Subzones) in pelagic “Ammonitico Rosso”
type facies as followed:
Chitinoidella Doben, 1963 – Chitinoidella boneti Doben,
Chitinoidella elongata Pop; Borziella Pop, 1997 – Borziella
slovenica (Borza);
Carpathella Pop, 1998 – Carpathella rumanica Pop;
Longicollaria Pop, 1997 – Longicolaria dobeni (Borza),
Longicolaria insueta (Řehánek);
Daciella Pop, 1998 – Daciella danubica Pop, Daciella
svinitensis Pop;
Dobeniella Pop, 1997 – Dobeniella tithonica (Borza), Do-
beniella cubensis (Furrazola-Bermudez), Dobeniella bermu-
dezi ( Furrazola-Bermudez) and
Popiella n.gen. – Popiella oblongata n.sp.
Further, three phylogenetic stocks were recognized in the
framework of vertical chitinoidellid distribution (Fig. 5):
1) The first one is derived from Longicollaria dobeni which
could have been the ancestor of larger crasicollarian loricas.
2) Lineage represented by Dobeniella tithonica gave rise to
forms having composed collars
3) Newly described Popiella oblongata could have been the
ancestor of Calpionellopsis and later Calpionellopsella type
forms.
Acknowledgments: The photographs were made by H. Brod-
nianska. The author is indebted to Prof. V. Houša (Prague),
Assoc. Prof. I. Lakova (Sofia), Prof. M. Mišík and Dr. J.
Michalík (Bratislava), for stimulating discussions and for crit-
ical reading of the manuscript. Technical support was provid-
ed by Mrs. E. Petriková. The author thanks also the Grant
Agency for Science, Slovakia (VEGA No. 2/2074/22) for fi-
nancial support.
References
Adatte T., Stinnesbeck W., Hubberten H. & Remane J. 1992: The
Jurassic-Cretaceous boundary in NE and Central Mexico – a
multistratigraphical approach. Actas de III Congreso Geológi-
co de Espana y VIII Congreso Latinoamericano de Geología 4,
23—29.
Altiner D. & Özkan S. 1991: Calpionellid zonation in north-western
Anatolia (Turkey) and calibration of the stratigraphic ranges of
some benthic foraminifers at the Jurassic-Cretaceous bound-
ary. Geol. Romana 27, 215—235.
Bakalova D. 1977: La succession a Calpionelles de la coupe pres du
village de Ginci, Bulgarie du Nord-Ouest. Compte Rendu, Ac-
adémie Bulgare de Sciences 30, 423—426.
Bonet F. 1956: Microfaunal zonation of the Cretaceous limestones
of the Eastern Mexico. Boletín de la Asociación Mexicana de
Geólogos Petroléros 8, 389—488 (in Spanish).
Borza K. 1965: Das Vorkommen der Gattung Chitinoidella Doben,
1962 im Oberjura der Westkarpaten. Geologický Sborník Slov-
enskej Akademie Vied 16, 3—5.
Borza K. 1966: Neue Arten der Gattung Chitinoidella Doben, 1962
in den Westkarpaten. Geologický sborník Slovenskej akademie
vied 17, 259—265.
Borza K. 1969: Die Mikrofacies und Mikrofossilien des Oberjuras
und der Unterkreide der Klippenzone der Westkarpaten. Slovak
Academy of Sciences Publishing House, Bratislava, 302.
Borza K. 1984: The Upper Jurassic-Lower Cretaceous parabiostrati-
graphic scale on the basis of Tintinninae, Cadosinidae, Stomi-
osphaeridae, Calcisphaerulidae and other microfossils from the
West Carpathians. Geol. Zbor. Geol. Carpath. 35, 5, 539—550.
Colom G. 1948: Fossil tintinnids: loricated Infusoria of the order of
the Oligotricha. J. Paleontology 22, 233—263.
Colom G. 1988: Tintinnids revision (Infusoria, Oligotricha, Lor-
icas). Species evolution in Paleozoic and Mesozoic marine
conditions. Rev. Españ. Paleontología 3, 71—104 (in Spanish).
Doben K. 1962: Paläontologisch-stratigraphische und fazielle Unter-
suchungen an der Jura/Kreide-Grenze in den bayrischen Kalkal-
pen zwischen Inn und salzach. Univ. München. Diss. 1—97.
Doben K. 1963: Ueber Calpionelliden an der Jura/Kreide – Gren-
ze. Mitt. Bayer. Staatsam. Paläontol. Hist. Geol. 3, 35—50.
Enay R. & Geyssant J.R. 1975: Faunes tithoniques des chaines bé-
tiques (Espagne méridionale). Mém. B.R.G.M. 86, 39—55.
Eshet Z. 1990: The palynostratigraphy of the Permian-Triassic
boundary in Israel: two approaches to biostratigraphy. Israel J.
Earth Sci. 39, 1—15.
Furrazola-Bermúdez G. 1965: New species of tintinnids from Up-
per Jurassic of Cuba. Instituto Cubano de Recursos Minerales,
Publ. Espec. 2, 1—29 (in Spanish).
Grandesso P. 1977: Tithonian praecalpionellids derived from Am-
monitico Rosso facies. Mem. Sci. Geol. 32, 1—14 (in Italian).
Lakova I. 1993: Middle Tithonian to Berriasian praecalpionellid
and calpionellid zonation of the Western Balkanides, Bulgaria.
Geol. Balcanica 23, 3—24.
Michalík J., Reháková D. & Halásová E. 1990: Stratigraphy of the
Jurassic-Cretaceous boundary beds in the Hlboč Valley
(Vysoká Unit of the Krížna Nappe, Malé Karpaty Mts). Knih.
ZPN 9a, 183—204.
Murgeaunu G. & Filipescu M.G. 1933: Calpionella carpathica n.sp.
dans les Carpathes roumaines. Notationes Biologicae 1, 63—64.
Olóriz F., Caracuel J.E., Marques B. & Rodríguez-Tovar F.J. 1995:
Tintinnid associations from Ammonitico Rosso of the Sierra
Norte (Mallorca). Rev. Españ. Paleontol. 77—93 (in Spanish).
Pop G. 1986. Calpionellids and correlation of Tithonian-Valangin-
ian formations. Acta Geol. Hung. 29, 93—102.
Pop G. 1989: Age and facies of the calpionellid formations from the
South Carpathians. In: Cretaceous of the Western Tethys. In:
Wiedmann J. (Ed.): Cretaceous of the Western Tethys. Pro-
ceedings of the 3rd International Cretaceous Symposium,
Tübingen 1987, 525— 542.
Pop G. 1997: Révision systématique des chitinoidelles tithoniennes
des Carpathes méridionales (Roumanie). C.R. Acad. Sci. Paris,
Sci. de la Terre Plan
ètes 324, 931—938.
Pop G. 1998a: Nouvelles chitino delles tithoniennes des Carpathes
méridionales (Roumanie). C.R.Acad.Sci. Paris, Sci. de la Terre
et Plan. 326, 817—822.
Pop G. 1998b: Stratigraphic distribution and biozonation of Titho-
nian Praecalpionellids and calpionellids from the South Car-
pathians. Rom. J. Stratigraphy 77, 4, 3—25.
CHITINOIDELLA TREJO, 1975 IN TITHONIAN PELAGIC SEQUENCES 379
Reháková D. 1995a: Upper Jurassic—Lower Cretaceous carbonate
microfacies and environmental models for the Western Car-
pathians and adjacent palaeogeographic units. Cretaceous Re-
search 16, 283—297.
Reháková D. 1995b: Calpionellid distribution in Upper Jurassic and
Lower Cretaceous sequences of the Western Carpathians. Min-
er. Slovaca 27, 308—318.
Reháková D. & Michalík J. 1992: Notes to ultrastructure study of
Upper Jurassic—Lower Cretaceous calpionellid tests. Acta
Univ. Carolinae Geol. 1—2, 107—110.
Reháková D. & Michalík J. 1993: Observation of ultrastructure of
the Upper Jurassic and Lower Cretaceous calpionellid tests.
Geol. Carpathica 44, 75—79.
Reháková D., Michalík J. & Ožvoldová L. 1996: New microbios-
tratigraphical data from several Lower Cretaceous pelagic se-
quences of Northern Calcareous Alps (preliminary results).
Mitt. Geol. Bergbaustud. Innsbruck 4, 57—81.
Reháková D. & Michalík J. 1997: Evolution and distribution of
calpionellids – the most characteristic constituent of Lower
Cretaceous Tethyan microplankton. Cretaceous Research 18,
493—504.
Reháková D. 2000a: Evolution and distribution of the Late Jurassic
and Early Cretaceous calcareous dinoflagellates recorded in
the Western Carpathian pelagic carbonate facies. Miner. Slova-
ca 32, 79—88.
Reháková D. 2000b: Calcareous dinoflagellate and calpionellid bio-
events versus sea-level fluctuations recorded in the West-Car-
pathian (Late Jurassic/Early Cretaceous) pelagic environments.
Geol. Carpathica 51, 4, 229—243.
Reháková D. 2002: Upper Jurassic and Lower Cretaceous micro-
granular calpionellids – implication for palaeoecology and
.palaeooceanography of the Western Carpathian pelagic envi-
ronments. ESSE WECA 2002, Abstract Book, 31—33.
Řehánek J. 1986: Chitinoidella insueta n.sp. (Protozoa incertae se-
dis) from the Tithonian of southern Moravia. Čas.
Mineral. Geol. 31, 3, 287—292.
Řehánek J. 1990: Calpionellids from the Jurassic and Cretaceous
limestones on the SE slope of the Bohemian Massif. Biostrati-
graphical and sedimentological study from Mesozoicum of the
Western Carpathian and Bohemian Massif. Knih. ZPN 9b,
185—208 (in Czech).
Remane J. 1969: Les possibilités actuelles pour une utilisation
stratigraphique des calpionelles (Protozoa incertae sedis, Cil-
iata?). Proceedings, Plankton Conference, Geneve 1967, 2,
559—573.
Remane J., Bakalova-Ivanova D., Borza K., Knauer J., Nagy I., Pop
G. & Tárdi-Filácz E. 1986: Agreement on the subdivision of
the standard calpionellid zones defined at the IInd Planktonic
Conference, Roma 1970. Acta Geol. Hung. 29, 5—14.
Skourtsis-Coroneou V. & Solakius N. 1999: Calpionellid zonation
at the Jurassic/Cretaceous boundary within the Vigla Lime-
stone Formation (Ionian Zone, western Greece) and carbon iso-
tope analysis. Cretaceous Research 20, 583—595.
Slipinski S.A. 1992: Larinotinae – a new subfamily of Trogossiti-
dae (Coleoptera), with notes on the construction of Trogossiti-
dae and related families of Cleroidea. Rév. Suisse Zool. 99, 2,
439—493.
Trejo M. 1975a: Mesozoic tintinnids of Mexico. Mém. Bureau
Réch. Géol. Min. 86, 95—104 (in Spanish).
Trejo M. 1975b: Mesozoic tintinnids of Mexico (taxonomy, age,
paleobiology). Bol. Asoc. Méxicane Geól. Petrol. XXVII, 10—
12, 329—449 (in Spanish).
Vašíček Z., Michalík J. & Reháková D. 1994: Early Cretaceous
stratigraphy, paleogeography and life in Western Carpathians.
Beringeria 10, 3—169.
Visscher H. 1970: On the occurrence of chitinoid loricas of Tintin-
nida in an Early Triassic palynological assemblage from King-
scourt, Ireland. Geological Survey of Ireland, Bull. 1, 61—64.
Visscher H. 1971: The Permian and Triassic of the Kingscourt outli-
er, Ireland – a palynological investigation related to regional
stratigraphical problems in the Permian and Triassic of West-
ern Europe. Geological Survey of Ireland, Spec. Pap. 1, 1—114.