GEOLOGICA CARPATHICA, APRIL 2005, 56, 2, 137147
www.geologicacarpathica.sk
Palynostratigraphy of the Maastrichtian dinosaur- and
mammal sites of the Râul Mare and Barbat Valleys
(Haþeg Basin, Romania)
JIMMY VAN ITTERBEECK
1*
, VALENTINA S. MARKEVICH
2
and VLAD CODREA
3
1
FWO-aspirant, Afdeling Historische Geologie, Katholieke Universiteit Leuven, B-3000 Leuven, Belgium;
jimmy.vanitterbeeck@geo.kuleuven.ac.be
*
Corresponding author: Tel. +32 16 326450
2
Institute of Biology and Pedology, Far East Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia;
markevich@ibss.dvo.ru
3
Catedra de Geologie-Paleontologie, Universitatea Babeº-Bolyai, Str. Kogãlniceanu 1, 3400 Cluj-Napoca, Romania; vcodrea@bioge.ubbcluj.ro
(Manuscript received February 3, 2004; accepted in revised form June 16, 2004)
Abstract: The palynomorph associations of the Upper Cretaceous dinosaur-bearing sediments of three different sites
(Pui, Toteºti-baraj and Nãlaþ-Vad) within the Haþeg Basin show a great similarity with the palynomorph associations of
the stratotypes of the Densuº-Ciula and Sânpetru Formations. The absence of volcanoclastic layers places the sediments
within the Sânpetru Formation. Palynological evidence indicates a Maastrichtian age of the vertebrate sites studied. The
age can be refined to the Early-Late Maastrichtian boundary interval based on the good correlation with the lower
Rogancian (France), the lower Garumnian (Spain) and the Gulpen Formation (Maastrichtian stratotype, The Nether-
lands). The ancient vegetation, dominated by ferns and bryophytes with disperse flowering plants and gymnosperm
trees, is indicative for a subtropical climate. Freshwater ferns flourished in the floodplain ponds. For the first time, the
megaspore genus Ghoshispora is recorded in Europe.
Key words: Maastrichtian, Southern Carpathians, Romania, Haþeg Basin, palynology.
Introduction
Uppermost Cretaceous (Campanian-Maastrichtian) vertebrate
assemblages are well known and documented from eastern
Asia and western North America. In Europe contemporary
faunas have been described from southern France, Spain and
Portugal (Le Loeuff 1991). With the exception of Laño (Pere-
da Suberbiola et al. 2000), the material collected at these lo-
calities is scarce and fragmentary and mammals appear to be
particularly rare, especially multituberculates, which are cur-
rently only known in Europe from the Haþeg Basin. The Upper
Cretaceous fauna recovered from the Haþeg Basin, Romania is
one of the most diverse and best preserved of these European
assemblages (Weishampel et al. 1991; Grigorescu et al. 1999).
In the present study, the palynomorph associations of the sedi-
ments at Toteºti-baraj, Nãlaþ-Vad, and Pui confirm the correla-
tion with the Sânpetru Formation and clarify the age of these
important vertebrate sites. The first Upper Cretaceous multitu-
berculate tooth (an incisor) in Europe was found in the Sânpet-
ru Formation in the Sibisel Valley (Grigorescu 1984), fol-
lowed by the first molars in the Barbat Valley at the Pui site
(Radulescu & Samson 1986; Grigorescu & Hahn 1987). The
sediments exposed at Pui are classically attributed to the Sân-
petru Formation (Grigorescu & Anastasiu 1990, p. 53). Re-
cently two new sites have been discovered in the Râul Mare
Valley (Fig. 1): Toteºti-baraj (Codrea et al. 2002) and Nãlaþ-
Vad (Smith et al. 2002). These sites have yielded numerous
well-preserved mammal teeth together with dinosaur egg nests
and dinosaur bones. On older geological maps of the Haþeg
Basin, exposures of Upper Cretaceous continental sediments
within the Râul Mare Valley are not indicated (Grigorescu
1983). On the basis of the regional geological framework, the
new Râul Mare sites have been tentatively attributed to the
Sânpetru Formation. The similar vertebrate assemblages seem
to confirm this hypothesis.
Stratigraphic setting of the Maastrichtian non-
marine deposits of the Haþeg Basin
Two Maastrichtian continental formations (Fig. 1) crop out
in the Haþeg Basin (Grigorescu & Anastasiu 1990), the Sân-
petru Formation (in the central-southern part) and the Densuº-
Ciula Formation (in the northwestern part). The latter is divid-
ed into three members and contains several volcanoclastic
intercalations, while the first is undivided and devoid of such
material. Both have yielded a rich dinosaur fauna. For the
Densuº-Ciula Formation, all the fossils have been recovered
from the middle member. The upper member is barren and
therefore its age is uncertain, although Grigorescu & Anasta-
siu (1990) and Grigorescu (1992) assumed a Paleogene age
based on the absence of dinosaur fossils. The lower part of this
formation rests unconformably on Upper Cretaceous marine
deposits. So the possible age of the lower and middle members
of the Densuº-Ciula Formation is limited to an interval between
the age of the underlying marine deposits and the extinction of
138 VAN ITTERBEECK, MARKEVICH and CODREA
the dinosaurs at the Cretaceous/Paleogene (K/P) boundary. The
lower contact of the Sânpetru Formation is not exposed and
therefore its stratigraphic position is more complicated. The
Sânpetru Formation and the middle member of the Densuº-Ciu-
la Formation have been correlated on the basis of their dinosaur
fauna and the age of both formations has always been treated to-
gether. Antonescu et al. (1983) described the freshwater gastro-
pods and palynomorph taxa (Table 2) recovered from the strato-
types of both units and this study provided further proof for the
correlation of the two stratigraphic units.
In the past, the marine sediments underlying the Densuº-Ciu-
la Formation have been estimated as rising up to the Early
Maastrichtian on the basis of foraminifer evidence (Grigorescu
& Anastasiu 1990; Pop et al. 1990). Taking into account the
Early Maastrichtian age of the underlying marine sediments and
the presence of Pseudopapilopollis praesubhercynicus in the
palynomorph associations (Antonescu et al. 1983), the dino-
saur-bearing deposits have previously been attributed to the
Late Maastrichtian (Grigorescu & Anastasiu 1990; Grigorescu
Fig. 1. a Localization of the Haþeg Basin (rectangle en-
larged in part b) within the regional geological framework.
b Simplified geological map of the Haþeg Basin with
the localization of the sites: Pui (A), Toteºti-baraj (B) and
Nãlaþ-Vad (C). 1 metamorphic formations (Late Prot-
erozoicCarboniferous) of the Danubian (b) Getic (c) and
Supragetic (a) Domains; 2 marine Cretaceous forma-
tions; 3 sedimentary pre-Cretaceous formations
(PermianJurassic); 4 post-Cretaceous sedimentary
formations (PaleogeneMiocene); 5 Densuº-Ciula For-
mation; 6 Sânpetru Formation. Black lines indicate ma-
jor faults. (Modified after M. Lupu in Grigorescu et al.
1990, 126.)
1992). Both arguments have proven unreliable and have been
refuted by Lopez-Martinez et al. (2001). Firstly, there is a con-
tinuous succession of palynomorph taxa across the K/P bound-
ary in the Tethys region, therefore ages should be based on as-
semblages rather than marker species. With the repositioning of
the Campanian-Maastrichtian boundary and new calibrations,
the foraminiferal data of the Haþeg Basin were in need of revi-
sion. Indeed, recently the underlying marine sediments have
been dated with calcareous nannoplankton as latest Campanian
(Grigorescu & Melinte 2001; Grigorescu & Csiki 2002). On the
basis of paleomagnetic data, Panaiotu & Panaiotu (2002) sug-
gested a Lower Maastrichtian age (Chron 31r, 68.771.0 Ma)
for the entire Sânpetru Formation.
Material and methods
During two different field campaigns, samples of organic
rich sediments were collected at the Pui (5 samples), Toteºti-
PALYNOSTRATIGRAPHY OF THE MAASTRICHTIAN DINOSAUR- AND MAMMAL SITES 139
baraj (10 samples) and Nãlaþ-Vad (6 samples) sites (Figs. 1, 2).
Nine samples in total have yielded palynomorphs; the other
samples were barren. The dark-coloured sediments at the Râul
Mare sites yielded rich palynomorph associations. The red-co-
loured sediments at the Pui site were all barren, only the
coarse-grained channel deposits, green in colour, have yielded
poor palynomorph associations.
Palynological techniques were applied to approximately
50 g of sediment per sample and involved treatment with cold
25% HCl, digestion for two days in HF (40%), followed by re-
peated hot baths (80 °C) in 25% HCl. The samples were
rinsed to neutrality between each step. No oxidation, heavy
liquid separation or ultrasonic treatment was applied. All resi-
dues were homogenized by stirring and sieved through nylon
filters with a mesh size of 15 µm. For sample NVD 7 rich in
megaspores, subsamples were sieved through 30 µm filters in
order to concentrate the megaspores. Both fractions were
stained with Safranin O and mounted with glycerine jelly.
Relative abundances were calculated from specimen counts
from the 15 µm slides. One or two slides from the 30 µm frac-
tion were scanned for rare species or especially well-preserved
specimens. Photographs were taken on a ZEISS Axioskop-2
light microscope equipped with a Sony DSC-S75 camera.
Results
Toteºti-baraj and Nãlaþ-Vad
The investigated samples have yielded rich, well-preserved
and diversified palynomorph associations (see Table 1 for a
counted list of taxa, Fig. 3). Spores dominate the associations
(7090 %), while gymnosperms and angiosperm pollen occur
in smaller quantities. Smooth trilete and monolete spores like
Leiotriletes, Cyathidites, Laevigatosporites dominate the as-
semblage. Osmundacidites, Polypodiaceoisporites and Glei-
cheniidites occur frequently. Bryophyte spores like Sphag-
numsporites and Selaginella are common. The most dominant
gymnosperm is Taxodiaceaepollenites hiatus, while bisaccate
pollen grains occur only in low numbers. The angiosperm as-
sociations consist of Normapolles (a.o. Nudopollis, Oculopol-
lis, Trudopollis) and Postnormapolles (a.o. Subtriporopolleni-
tes, Triatriopollenites) type of pollen grains. The angiosperm
pollen are very diverse without a true dominance of one of the
taxa. One of the samples (Table 1, NVD 7) has yielded an in-
teresting assemblage of megaspores of Ghoshispora and Azol-
la (Fig. 4). In spite of their similar morphological characteris-
tics, the Ghoshispora specimens show a distinct difference in
size. They can be divided into a large (ca. 125 µm) and a small
(ca. 50 µm) form (Fig. 4). Both forms have been attributed to
Ghoshispora bella, as the size of this species is highly variable
(Srivastava 1971).
Pui site
Only one sample from green channel deposits was produc-
tive and yielded a very poor palynological assemblage (Ta-
ble 1), both in number of taxa and in number of specimens.
Only the taxa with a relatively high abundance rate in the oth-
er samples have been observed at the Pui site. In spite of the
low number of counted specimens (Table 1), the assemblage
shows the same trends as for the other sites. Spores dominate
the assemblage, gymnosperm pollen occur relatively frequent-
ly while angiosperm pollen are rare.
Fig. 2. Schematic view of the exposures at Pui (A), Toteºti-baraj (B) and Nãlaþ-Vad (C) with the location of the palynological samples
(associations given in Table 1), black pentagons and ellipses indicate the vertebrate fossils.
140 VAN ITTERBEECK, MARKEVICH and CODREA
Pui/3
NVD/1
NVD/7
NVD/11 TOT/7
TOT/8 TOT/10 TOT/18 TOT/19
Spores
22 56.4 204 71.8 233 74.9 187 74.5 18 13.6 190 72.5 71 62.8 316 90.3 444 89.2
Acanthotriletes sp.
2 0.4
Adiantum mirum Chlonova, 1960
1 0.4
Anapiculatisporites dawsonensis Reiser et Williams, 1969
*
1 0.4
Appendicisporites sp.
2 0.4
Appendicisporites tricornitatus Weyland et Greifeld, 1953
1 0.4
3 0.6
Azolla sp.
29 9.3
Baculatisporites comaumensis (Cookson) Potonié, 1956
1 0.2
Balmeisporites sp.
1 0.2
Ceratosporites equalis Cookson et Dettmann, 1958
3 1.1
Cicatricosisporites dorogensis Potonié et Gelletich, 1933
*
5 1.8 4 1.3 6 2.4
11 4.2 1 0.9 47 13.4 34 6.8
Cicatricosisporites furcatus Deak, 1963
2 0.8
3 0.6
Cicatricosisporites sp.
1 0.4
Cicatricosisporites subrotundus Brenner, 1963
*
1 0.4
Cingutriletes cf. C. clavus (Balme) Dettmann, 1963
1 0.4
Chomotriletes fragilis Pocock, 1962
3 1.1 2 0.6 1 0.4
2 0.8
3 0.9 3 0.6
Chomotriletes sp.
7 2.5
1 0.4
Concavisporites junctus (Kara-Murza) Semenova, 1970
1 0.3
Concavissmisporites asper (Bolchovitina) Pocock, 1964
3 1.1
1 0.4
3 0.9
Converrucosisporites proxigranulatus Brenner, 1963
*
1 0.4
Cyathidites australis Couper, 1953
3 1.1 3 1.0 7 2.8
20 7.6 5 4.4 8 2.3 10 2.0
Cyathidites minor Couper, 1953
4 10.3 36 12.7 21 6.8 56 22.3 6 4.5 14 5.3 22 19.5 110 31.4 100 20.1
Densoisporites perinatus Couper, 1958
*
2 0.6
Densoisporites velatus Weyland et Krieger, 1953
5 1.8
1 0.9 3 0.9 5 1.0
Dyctiophyllidites harrisii Couper 1953
1 0.3 1 0.2
Echinatisporites levidensis (Balme) Srivastava, 1972
2 0.6
Foraminisporis asymmetricus (Cookson et Dettmann) Dettmann, 1963
1 0.2
Foveosporites canalis Balmé, 1957
1 0.2
Foveosporites cenomanicus (Chlonova) Schvetzova, 1976
1 0.3 1 0.4
Foveosporites sp.
2 0.6
Gabonisporites labyrintus Srivavastava, 1972
3 1.1 5 1.6
11 4.2
Ghoshispora bella (Kondinskaya) Srivastava, 1978
29 9.3
Gleicheniidites circinidites (Cookson) Dettmann, 1963
*
8 2.8 4 1.3
2 0.8
1 0.3 1 0.2
Gleicheniidites dicarpoides Grigoryeva, 1961
4 1.4
Gleicheniidites laetus (Bolchovitina) Bolchovitina, 1968
13 4.6 5 1.6 1 0.4
6 2.3
2 0.4
Gleicheniidites rasilis (Bolchovitina) Bolchovitina, 1968
1 0.4
Gleicheniidites senonicus Ross,1949
6 2.1
2 1.8
Gleicheniidites sp.
1 2.6
2 0.8
1 0.3
Gleicheniidites triplex (Bolchovitina) Krutzsch, 1959
2 0.8
1 0.3
Hamulatisporites rugulatus (Couper) Srivastava, 1972
1 0.4
2 0.6
Heliosporites kemensis (Chlonova) Srivastava
4 1.4 8 2.6
4 1.1 1 0.2
Intrapunctatosporis sp.
1 2.6
Klukisporites notabilis Srivastava, 1972
13 4.2
1 0.4
Klukisporites variegatus Couper, 1958
2 0.7
6 1.7 1 0.2
Klukisporites visibilis (Bolchovitina) Pocock, 1962
4 1.6
Kuylisporites lunaris Cookson et Dettmann, 1958
1 0.3
Laevigatosporites major (Verbitskaja) Zhang Q.B., 1984
1 0.2
Laevigatosporites ovatus Willson et Webster, 1946
5 12.8 22 7.7 20 6.4 24 9.6 2 1.5 4 1.5 16 14.2 38 10.9 100 20.1
Laevigatosporites ovoideus Takahashi, 1961
10 3.5
3 0.9
Leiotriletes rotundiformis Bolchovitina, 1956
1 0.3 1 0.2
Leiotriletes sp.
6 15.4 33 11.6 34 10.9 57 22.7 10 7.6 52 19.8 20 17.7 37 10.6 89 17.9
Leptolepidites bullatus (Van Hoeken- Klinkenberg) Srivastava, 1972
2 0.6
Leptolepidites verrucatus Couper, 1953
10 3.5 7 2.3 1 0.4
5 1.9
6 1.2
Liburnisporites adnacus Srivastava, 1972
1 0.4
5 1.9
4 0.8
Matonisporites equiexinus Couper, 1958
1 0.3
Matonisporites sp.
2 0.4
Neoraistrickia sp.
1 0.2
Neoraistrickia truncata (Cookson) Potonié, 1956
*
2 0.6
Nevesisporites radiatus (Chlonova) Srivastava, 1972
1 0.3
2 0.6 1 0.2
Ophioglossum sp.
1 0.4
Osmundacidites sp.
4 1.1 11 2.2
Osmundacidites wellmanii Couper, 1953
1 2.6 5 1.8 3 1.0 6 2.4
16 6.1
9 2.6 9 1.8
Polycingulatisporites reduncus (Bolchovitina) Playford et Dettmann, 1965
1 0.4 7 2.3
1 0.4
Polypodiaceoisporites cyclocingulatus Krutzsch, 1967
1 0.4
6 2.4
Polypodiaceoisporites potoniei Kedves, 1961
1 0.2
Polypodiaceoisporites variabilis Pacltova, 1970
4 10.3
15 5.7 3 2.7 9 2.6 23
Polypodiidites speciosus (Harris) Archangelsky, 1972
1 0.3
Retitriletes rotundiformis (Kara-Murza) Krutzsch, 1963
3 1.0
2 0.6 5 1.0
Rouseisporites reticulatus Pocock, 1962
1 0.4 5 1.6 2 0.8
2 0.8
4 0.8
Rouseisporites sp.
1 0.2
Schizaeoisporites cicatricos Sole de Porta, 1972
*
5 2.0
10 2.9 2 0.4
Selaginella intertexta Krasnova, 1961
1 0.4
Selaginella sp.
15 5 .7
Sphagnumsporites antiquasporites (Wilson et Webster) Potonié, 1956
2 0.4
Sphagnumsporites sp.
5 1.8 3 1.0
1 0.9 1 0.3
Sphagnumsporites psilatus (Ross) Couper, 1957
1 0.4
Spinatisporites sp.
1 0.4
8 1.6
Todisporites minor Couper, 1958
1 0.2
Todisporites sp.
1 0.4
Triletes bettianus Srivastava, 1972
*
2 0.7 5 1.6
Tuberosisporites sp.
*
2 0.7 17 5.5
2 0.8
Table 1: Palynomorph associations (samples indicated on Fig. 2, * indicates the species in common with the Gulpen Formation (accord-
ing to Kedves & Herngreen 1980).
PALYNOSTRATIGRAPHY OF THE MAASTRICHTIAN DINOSAUR- AND MAMMAL SITES 141
Table 1: Continuing.
Pui/3
NVD/1
NVD/7
NVD/11
TOT/7
TOT/8
TOT/10 TOT/18 TOT/19
Gymnosperms
12 30.8 40 14.1 39 12.5 53 21.1 47 35.6 62 23.7 18 15.9 17 4.9 37 7.4
Abiespollenites editus (Chlonova) Chlonova, 1976
1 2.6
Abietinaepollenites sp.
2 1.5
Alisporites aequalis Mädler, 1964
2 5.1 3 1.1
Alisporites bilateralis Rouse, 1959
*
4 1.4 6 1.9 5 2.0
1 0.3
Alisporites bisaccus Rouse, 1959
1 0.2
Araucariacites sp.
3 1.1 1 0.3 3 1.2
1 0.2
Cedripites parvisaccatus (Sauer) Krutzsch, 1971
2 0.7 3 1.0 5 2.0 6 4.5
1 0.9
Cedruspollenites sp.
1 2.6
Classopollis classoides (Pflug) Pocock et Jansonius, 1961
*
1 0.3 1 0.4
5 1.9
3 0.9
Classopollis sp.
1 2.6
3 0.6
Ginkgocycadophytus sp.
2 5.1 5 1.8 4 1.3 4 1.6
3 1.1 5 4.4 2 0.6 3 0.6
Inaperturopollenites sp.
1 0.4
Piceapollenites sp.
21 15.9
5 4.4
1
Pinuspollenites sp.
5 12.8 3 1.1 14 4.5 9 3.6 12 9.1 15 5.7 2 1.8 2 0.6 12 2.4
Podocarpidites ellipticus Cookson, 1947
1 0.3
2 0.8
Podocarpidites multessimus (Bolchovitina) Pocock, 1962
*
5
1.8
Taxodiaceaepollenites hiatus (Potonié) Kremp, 1949
15 5.3 9 2.9 25 10.0 6 4.5 37 14.1 5 4.4 9 2.6 16 3.2
Pui/3
NVD/1
NVD/7
NVD/11
TOT/7
TOT/8
TOT/10 TOT/18 TOT/19
Angiosperms
5 12.8 40 14.1 39 12.5 11 4.4 67 50.8 10 3.8 24 21.2 17 4.9 17 3.4
Atlantopollenites choffatii Diniz, Kedves et Simoncsics, 1977
1 0.4
Coryluspollenites sp.
12 9.1 0.0 2 1.8
1 0.2
Echimonocolpites minutus Salard-Cheboldaeff, 1978
1 0.4 3 1.0
2 0.8
Elytranthe striatus Couper, 1953
1 2.6
1 0.2
Hofkeripollenites capsula (Pflug) Kedves et Hergreen, 1980
*
5 4.4 7 2.0 2 0.4
Hofkeripollenites triangulus Kedves et Hergreen, 1980
*
1 0.4
Interpollis microsupplingensis Krutzsch, 1961
1 0.8
Interpollis supplingensis (Pflug) Krutzsch, 1961
*
1 0.4 3 1.0
1 0.9
Interpollis cf. velum Krutzsch, 1961
*
1 0.8
Interporopollenites sp.
5 1.8 11 3.5 1 0.4
2 0.6
Liliacidites sp.
1 0.9
Maestrichtipollenites concavus Kedves et Hergreen, 1980
*
1 0.4
Minorpollis maestrichtiensis Kedves et Hergreen, 1980
*
1 0.3
1 0.3
Nudopollis apertus (Pflug) Pflug, 1953
*
1 0.4
Nudopollis minutus Zaklinskaja, 1963
*
5 1.6
1 0.3
Nudopollis terminalis (Thomson et Pflug) Pflug, 1953
*
2 0.7
Oculopollis minoris Krutzsch, 1973
*
2 0.7
5 1.9
1 0.3
Oculopollis sp.
1 0.3 2 0.4
Platycarya sp.
1 0.3 1 0.2
Plicapollis conserta Pflug, 1953
*
2 0.7 1 0.3
Plicapollis serta Pflug, 1953
*
1 0.3
3 0.6
Plicapollis cf. silicatus Pflug, 1953
*
1 0.4
Pompeckjoidaepollenites platoides (Pflug, 1953) Krutzsch, 1967
1 0.3
Portniaginaepollenites maestrichtiensis Kedves et Hergreen, 1980
*
1 0.8
Portniaginaepollenites minor Kedves et Hergreen, 1980
*
1 0.8
Quercites sparsus ( Martynova) Samoilovich, 1961
4 1.6
Retitricolpites georgensis Brenner, 1963
1 2.6
Retitricolpites sp.
1 0.4
Subtriporopollenites constans Pflug, 1953
*
2 0.6
2 1.5
Subtrudopollis sp.
2 0.6
Subtrudopollis subtrudens (Pflug) Krutzsch, 1967
*
2 0.6
Thomsonipollis magnificus (Thomson et Pflug) Krutzsch, 1960
1 0.2
Triatriopollenites aroboratus Pflug, 1953
10 3.5
Triatriopollenites bituites (Potonié) Thomson et Pflug, 1953
*
4 1.4
30 22.7
Triatriopollenites confusus Zaklinskaya, 1963
2 1.5
1 0.9
1 0.2
Triatriopollenites lubomirovae ( Gladkova) Kedves, 1974
*
5 1.8 1 0.3
14 10.6
Triatriopollenites roboratus Pflug, 1953
* 1 2.6 1 0.4
Tricolpites hians Stanley, 1965
1 0.4
2 1.8
Tricolpites microreticulatus Boltenhagen et Potonié, 1965
1 2.6
Tricolpites reticulatus Cookson, 1947
1 0.2
Tricolpites sp.
1 2.6 1 0.4 3 1.0 1 0.4
1 0.3 1 0.2
Tricolpites vulgaris (Pierce) Srivastava, 1969
1 0.4
Tricolporopollenites sp.
1 0.4
Triporopollenites cretacicus Kedves et Herngreen, 1980
*
1 0.3
1 0.8
Triporopollenites sp.
1 0.4 2 1.5
1 0.9 1 0.3
Trudopollis maestrichtiensis Kedves et Herngreen, 1980
*
3 1.1
2 0.4
Trudopollis cf. parvotrudens Pflüg, 1953
*
1 0.4
Trudopollis sp.
3 1.0
10 8.8
Ulmoideipites tricostatus Anderson, 1960
1 0.9
Vacuopollis sp.
*
1 0.2
Total
39 100.0 284 100.0 311 100.0 251 100.0 132 100.0 262 100.0 113 100.0 350 100.0 498 100.0
142 VAN ITTERBEECK, MARKEVICH and CODREA
Interpretation
Correlation
Due to the lack of quantative data, it is difficult to compare
the list (Table 2) of Antonescu et al. (1983) with the data of
the present paper. These authors describe the associations
from the stratotypes of the Sânpetru and Densuº-Ciula Forma-
tions as follows: smooth trilete microspores of the type Leiot-
riletes and Dictyophyllidites are dominant, followed by spores
with a cingulum like Polypodiaceoisporites, disaccate pollen
are nearly absent while Inaperturopollenites are frequent, pol-
len of the Normapolles and Postnormapolles group form the
Fig. 3. 1 Hofkeripollis capsula (Toteºti, slide 18/1). 2 Nudopollis minutus (Toteºti, slide18/1). 3 Oculopollis minoris (Toteºti, slide
18/1). 4 Minorpollis maestrichtiensis (Toteºti, slide 18/1). 5 Retitricolpites sp. (Nãlaþ-Vad, slide 11/1). 6 Quercites sparsus (Nãlaþ-
Vad, slide 11/1). 7 Tricolpites vulgaris (Nãlaþ-Vad, slide 11/1). 8 Trudopollis cf. parvotrudens (Nãlaþ-Vad, slide 11/1). 9 Triatri-
opollenites confusus (Toteºti, slide 19/2). 10 Cyathidites australis (Toteºti, slide 19/3). 11 C. australis (Toteºti, slide 19/1). 12
Cyathidites minor (Toteºti, slide 19/1). 13 C. minor (Nãlaþ-Vad, slide 11/1). 14 Leiotriletes rotundiformis (Toteºti, slide19/3). 15
Laevigatisporites ovatus (Nãlaþ-Vad, slide 11/1). 16 Polypodiaceoisporites variabilis (Toteºti, slide 18/1). 17 Gleicheniidites triplex
(Toteºti, slide 18/1). 18 G. circinidites (Toteºti, slide 18/1). 19 Retitriletes rotundiformis (Toteºti, slide 18/1). 20 Rouseisporites re-
ticulatus (Toteºti, slide 19/3). 21 Polypodiidites speciosus (Toteºti, slide 18/1). 22 Echinatisporites levidensis (Toteºti, slide 18/1).
23 Sphagnumsporites antiquasporites (Toteºti, slide 19/2). 24 Heliosporites kemensis (Toteºti, slide 19/3). 25 Cicatricosisporites
dorogensis (Toteºti, slide 19/3). 26 Schizaeoisporites cicatricos (Toteºti, slide 18/1). 27 Chomotriletes fragilis (Toteºti, slide 18/1).
(Figs.19 scale bar = 10 µm; Figs.1027 scale bar = 20 µm).
foundation of the association. These characteristics apply to
the palynomorph associations of the different sites (Table 1,
see previous section). Cicatricosisporites occurs frequently
while Antonescu et al. (1983) indicate that these types of
spores are rare in the stratotype sections. Generally the an-
giosperm taxa observed in this study are very similar to those
recognized by Antonescu et al. (1983), although Pseudo-
papilopollis praesubhercynicus, considered by the same au-
thors as an important marker, has not been observed in the
studied samples. In spite of these differences, the palynomor-
ph associations allow the correlation of the three sites with the
middle member of the Densuº-Ciula Formations and Sânpetru
Formation. The absence of volcaniclastic material clearly
places the sediments studied within the Sânpetru Formation.
PALYNOSTRATIGRAPHY OF THE MAASTRICHTIAN DINOSAUR- AND MAMMAL SITES 143
Fig. 4. Megaspores (Nãlaþ-Vad). 12 Azolla sp. with detail of glochidia, SEM. 34 Azolla sp. 5 Ghoshispora bella (large), SEM.
6 Ghoshispora bella (large). 7 Ghoshispora bella (small), SEM. 89 Ghoshispora bella (small).
Biostratigraphy
The extant genus Azolla is known in the fossil record since
the Late Cretaceous. From this period, the genus is well-
known from Campanian and especially Maastrichtian strata of
North America (Batten & Kovach 1989, 1990).
Ghoshispora bella has been recorded from Senonian-Dani-
an deposits (Kondinskaya 1966) and the Maastrichtian of
North America: from the Whitemud Formation, Alberta, Can-
ada (Srivastava & Binda 1969), the Edmonton and the Mary
River Formation, Alberta, Canada (Snead 1969), the Hell
Creek Formation, Montana, U.S.A. (Norton & Hall 1967) and
the Lance Formation, Wyoming (Farabee & Canright 1986).
Although Srivastava (1971) considers G. bella a valuable
stratigraphic marker for Maastrichtian strata, Gunther & Hills
(1972) observed it throughout the Brazeau Formation (Cam-
panian-Maastrichtian), Alberta, Canada and Speelman & Hills
(1980) observed it in the Pakowki, Foremost and Oldman For-
mations (Santonian-Campanian), Alberta, Canada; therefore
the stratigraphic range of this species has to be extended into
the Santonian. The megaspore genus Ghoshispora has never
been recorded in Europe before.
The presence of the genera Interpollis and Nudopollis clear-
ly place the studied dinosaur-bearing strata within the Maas-
trichtian (Goczan et al. 1967).
The spore assemblage, rich in taxa and dominated by
Cyatheaceae, Gleicheniaceae, Schizaceae, resembles the pa-
lynozone A of Ashraf & Erben (1986), which occurs in the
lower Rognacian and the lower Garumnian (regional stages of
the Early Maastrichtian). With the new positioning of the
CampanianMaastrichtian boundary (Odin & Lamaurelle
2001), the age of the lower Rognacian has been revised as
Late Campanian-Early Maastrichtian (Garcia & Vianey-Liaud
2001; Cojan et al. 2003).
Portniaghina (1971, 1973, 1981) established a palynologi-
cal zonation for the flysh deposits of the Skale Zone of the
Carpathians. Campanian and Early Maastrichtian palynozones
are characterized by abundant Normapolles grains. Campa-
nian associations are dominated by forms with long annuli and
oculi. The Early Maastrichtian associations are characterized
by the absence of typical Campanian taxa like Lusatipollis and
Longanulipollis, by the presence of Convexipollis and by a va-
riety of species of Interporopollenites, Plicapollis, Pseudo-
papilopollis and Trudopollis. Upper Maastrichtian deposits
144 VAN ITTERBEECK, MARKEVICH and CODREA
Sibiºel
Convexipollis cf. convexigerminalis Krutzsch, 1967
Oculopollis cf. baculotrudens (Pflug) Zaklinskaya, 1963 cf.
Oculopollis sibiricus Zaklinskaya, 1963
Papilopollis? spp.
Plicapollis cf. conserta Pflug, 1953
Pseudopapillopollis praesubhercynicus (Goczan) Goczan, 1967
Toteºti
Callidasporites sp.
Carnisporites granulatus Schulz
Cyathidites minor Couper, 1953
Dictyophyllidites equiexinus (Couper) Dettmann, 1963
Leiotriletes maxoides Krutzsch
Plicifera delicata (Bochovitina) Bolchovitina, 1968
Polycingulatispotites reduncus (Bolchovitina) Playford et Dettmann, 1965
Triporoletes asper Srivastava, 1972
Oculopollis semimaximus Krutzsch
Oculopollis orbicularis Goczan, 1964
Plicapollis serta Pflug, 1953
Suemeghipollis triangularis Goczan, 1964
Concentricystes sp.
Deflandrea sp.
Ciula
Azolla sp.
Cicatricosisporites spp. (rare)
Corrugatisporites toratus Weyland et Greifeld, 1953
Dictyophyllidites spp.
Echinatisporites longechinus Krutzsch, 1959
Laevigatosporites spp.
Leiotriletes spp.
Polypodiisporites spp.
Polypodoaceoisporites spp.
Spore type B
Triletes sp.
Triporoletes cf. tornatilis Srivastava, 1972
Triporoletes spp.
Zlivisporites blanensis Pacltova, 1961
Inaperturopollenites sp.
cf. Comptonia sp.
cf. Engelhardtioidites sp.
Interporopollenites cf. gracilis Krutzsch, 1960
Interporopollenites proporus Weyland et Greifeld, 1953
Liliacidites sp.
cf. Minorpollis sp.
Oculopollis cf. baculotrudens (Pflug) Zaklinskaya, 1963
Oculopollis cf. orbicularis Goczan, 1964
Papilopollis spp.
Pistillipollenites sp.
Plicapollis cf. conserta Pflug, 1953
Plicapollis cf. pseudoexcelsus (Krutzsch) Krutzsch, 1961
Pseudoculopollis sp.
Pseudopapilopollis praesubhercynicus (Goczan) Goczan, 1967
Pseudoplicapollis peneserta (Pflug) Krutzsch, 1967
Semioculopollis praedicatus (Weyland et Krieger) Krutzsch, 1967
Spermatites spp.
Subtriporopollenites aff. constans Pflug, 1953 subsp. 1 & 2
Suemeghipollis triangularis Goczan, 1964
Tricolpites sp.1 & 2
Tricolporites sp.
Tricolporopollenites spp.
Trudopollis ex gr. imperfectus (Pflug) Pflug, 1953
Trudopollis? spp.
Vacuopollis sp.
Table 2: Palynomorph taxa from the stratotypes of the middle mem-
ber of the Densuº-Ciula Fomation (Ciula locality), of the Sânpetru
Formation (Sibiºel locality) (Antonescu et al. 1983) and Toteºti
borehole (Stancu et al. 1980).
have yielded associations where species of Extratripotopolle-
nites, Subtriporopollenites, Tetrapollis and Trudopollis are
more abundant and older Normapolles taxa like Quedlinburgi-
pollis and some species of Oculopollis are absent. On the basis
of these characteristics, the continental sediments of the Haþeg
Basin show a good correlation with the Maastrichtian strata of
the flysch deposits of the Skale Zone of the Carpathians.
Antonescu et al. (1983) used Pseudopapilopollis praesub-
hercynicus to correlate the strata of the Haþeg Basin with the
Upper Maastrichtian strata of Hungary. The absence of this
taxon makes the correlation with the Senonian sediments of
Hungary less clear. According to Goczan & Siegl-Farkas
(1990), Subtriporopollenites constans is a characteristic spe-
cies of the Hungarian Upper Maastrichtian and therefore, in
spite of the absence of P. praesubhercynicus in our samples,
the correlation with the Palaeostomocystis bakonyensis-
Pseudopapillopollis praesubhercynicus Assemblage Zone,
characteristic for the Hungarian Upper Maastrichtian, is main-
tained. In the new integrated biostratigraphy for the Romanian
Cretaceous (Ion et al. 1998), Antonescu considers P. praesub-
hercynicus as the marker species of the uppermost Lower
Campanian to the lowermost Lower Maastrichtian. Due to the
absence of any of the marker-species, the associations studied
(Table1) could not be fitted into the zonation as proposed by
Ion et al. (1998). Further comparison with Romanian palyno-
morph associations of Maastrichtian age is impossible due to
the lack of published data (Antonescu 2000, p. 9).
The palynological assemblage of the Maastrichtian strato-
type (Kedves & Herngreen 1980), more precisely of the Lixhe
and Lanhaye Members of the Gulpen Formation is very simi-
lar to the associations of the Romanian sites. Both associations
have 23 angiosperm taxa in common on the species level (in-
dicated with * in Table 1). Both members have been dated as
early Late Maastrichtian (Jagt 2002).
On the basis of the arguments given above, the dinosaur-
bearing sediments are considered to be Maastrichtian in age.
The good correlation with the early Rognacian and the Lixhe
Member of the Gulpen Formation, shows that the age of the
three analysed sections can probably be refined to the Early
Late Maastrichtian boundary interval. The palynological evi-
dence presented here corroborates the interpretation of the pa-
leomagnetic data of Panaiotu & Panaiotu (2002), who place
the Sânpetru Formation within the 31r-magnetochron (68.7
71.0 Ma).
Paleoenvironment
As stated earlier, the palynomorph association of the sites
studied shows a good correlation with palynozone A of the
lower Rognacian (Ashraf & Erben 1986). On the basis of the
botanical affinities of the observed taxa, these authors demon-
strated a tropical to subtropical evidence for this palynozone.
The same line of reasoning can be applied on the dinosaur-
bearing sediments of the Haþeg Basin. The presence of cal-
crete paleosols has been demonstrated for the sections studied
in the present paper by Van Itterbeeck et al. (2004) and for
other sections in the Haþeg Basin by Grigorescu & Anastasiu
(1990) and Grigorescu & Csiki (2002). According to Mack &
James (1994), these type of paleosols are characteristic of the
PALYNOSTRATIGRAPHY OF THE MAASTRICHTIAN DINOSAUR- AND MAMMAL SITES 145
Fig. 5. Summary diagram with the Campanian and Maastrichtian
geochronology and magnetostratigraphy (Gradstein et al. 1995) with
the age and magnetostratigraphy of the Rognacian (Galbrun 1997;
Garcia & Vianey-Liaud 2001), the age of the Lixhe and Lanhaye
members (Jagt 2002), the magnetostratigraphy of the Sânpetru For-
mation (Panaiotu & Panaiotu 2002) and the estimated age of the
Barbat and Râul Mare sites.
dry subtropical climatic zone. Thus the climate, during the
deposition of the dinosaur-bearing sediments of the Haþeg Ba-
sin, is interpreted as subtropical.
This climatic interpretation is supported by the presence of
the Azolla and Ghoshispora megaspores. Azolla is a free float-
ing water fern in freshwater ponds and lakes in a perennially
to seasonally wet tropical to warm-temperate habitats (Yi et al.
2003). Ghoshispora bella, also a freshwater fern (Kondin-
skaya 1966), is frequently found together with Azolla (Srivas-
tava 1971) and thus indicative of similar ecological condi-
tions. These megaspores together with freshwater algae like
Chomotriletes (= Concentricystes) clearly prove that the sedi-
ments of sample NVD7 were deposited in a freshwater pond.
Indeed, based on sedimentology, these sediments have been
interpreted as abandoned channel deposits (Van Itterbeeck et
al. 2004).
Exposures of Upper Cretaceous continental sediments were
not reported in the Râul Mare Valley prior to the discovery of
the Toteºti-baraj and Nãlaþ-Vad sites (Codrea et al. 2002;
Smith et al. 2002). They were reported, however, from the
subsurface (Stancu et al. 1980) with data on their palynologi-
cal content (Table 2). The dinoflagellate cyst Deflandrea sp. is
mentioned from the borehole sediments but has not been
found in the outcrop samples. This type of cyst is known from
lagoonal, estuarine and brackish water sediments (Powell et
al. 1996, p. 170; Stover et al. 1996, p. 716) and therefore the
continental environments of the dinosaur sites were located
close to the ancient shoreline.
Conclusions
The palynomorph associations of three important vertebrate
sites (Pui, Toteºti-baraj and Nãlaþ-Vad) perfectly match the
characteristics of the palynology of the stratotypes of the
Densuº-Ciula and Sânpetru Formations as given by Antones-
cu et al. (1983). The absence of volcanoclastic layers typical
of the Densuº-Ciula Formation, clearly places the studied sed-
iments within the Sânpetru Formation.
The associations show a good correlation with palynozone
A as described by Ashraf & Erben (1986) from the lower Ro-
gnacian (France) and lower Garumnian (Spain) (Late Campa-
nianEarly Maastrichtian). There is also a good correlation
with the palynomorph associations of the Maastrichtian strato-
type (Kedves & Herngreen 1980). On the basis of these corre-
lations and the general taxonomic composition of the associa-
tions (Fig. 5), the age of the vertebrate sites from the Râul
Mare and Barbat Valleys is estimated as Maastrichtian and can
be refined to the Early/Late Maastrichtian boundary interval.
The palynological data corroborates the age estimations for
the Sânpetru Formation of Panaiotu & Panaiotu (2002).
According to the palynology, the ancient vegetation can be
characterized as an open vegetation dominated by ferns and
bryophytes with some disperse flowering plants and a few
gymnosperm trees under a subtropical paleoclimate. In the
floodplain ponds, freshwater ferns with megaspores like Azol-
la and Ghoshispora flourished.
Acknowledgments: We greatly appreciate the time and effort
of all the participants in the field: Cristina Fãrcaº, Steffi M.,
Paul Grovu, Paul Dica, Virgil Benedek, Cladiu Chendeº,
Sergiu Hossu, Emanoil and Liana Sãsaran, Suzanne Watrin,
Géraldine Garcia, Stijn Goolaerts, Pascal Godefroit, and Thi-
erry Smith. JVI is a Research Assistant of the Fund for Scien-
tific Research Flanders, Belgium (FWO Vlaanderen)
and thanks his supervisors, Prof. Noel Vandenberghe and
Prof. P. Bultynck whose constructive comments improved the
paper significantly. The fieldwork was supported in particular
by travel grants from the Dirk Vogel Fonds KUL to JVI.
Field vehicles were kindly provided by Fabricom NV (2001)
and by Ford NV (2002). Financial support for this project has
been provided by Research Project MO/38/004 of the Belgian
Federal Office for Scientific, Technical and Cultural Affairs
(DWTC-SSTC) and Grants 04-1-P25-053 Origin and evolu-
tion of biosphere and 04-1-06-002 Groundlines of biodiver-
sity preservation in Russia of the Programme of the Presidi-
um of the Russian Academy of Sciences.
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