the budurone microvertebrate site from the maastrichtian of the

ACTA PALAEONTOLOGICA ROMANIAE V. 6 (2008), P. 49-66.
THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE
HAŢEG BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
Zoltán CSIKI1, Ana IONESCU2 & Dan GRIGORESCU3
Abstract: Microvertebrate fossil sites contain mixed remains of different aquatic and terrestrial taxa,
characterized by the small size of the skeletal elements, dominance of the resistant skeletal parts, high
degree of disarticulation and extensive taphonomic modifications. Microvertebrate sites are useful for
paleoecological reconstructions in continental settings since they potentially sample a broad range of
microhabitats due to their mode of genesis. Several microvertebrate sites were discovered in the Upper
Cretaceous (Maastrichtian) continental deposits of the Haţeg Basin (southwestern Carpathians) beginning
with 1983. Some of the most important sites are located around Vălioara village, in the northwestern part of
the Haţeg Basin. One of these is the Budurone site, remarkable especially for the association of the
vertebrates with plant remains, in a dark grey mudstone. Sedimentological study of the locality suggests
the sediments were deposited within a small floodplain pond or marsh. Diverse seed and palynomorph
assemblages were recovered from the site; the seeds are referred to indeterminate angiosperms, while
polypodiacean fern spores and especially Normapolles pollen dominate the palynological assemblage. The
plant assemblage allows the reconstruction of the vegetation around the pond as being made up of a
herbaceous angiosperm-fern ground cover and a thick low angiosperm canopy. The local food web can
also be reconstructed, being dominated by small-sized aquatic and semi-aquatic taxa.
Keywords: Haţeg Basin, Maastrichtian, microvertebrates, palynomorphs, paleoecology, paleoenvironment
INTRODUCTION
The European Late Cretaceous continental
assemblages were (and are continuing to be)
less well known than their Asian, South
American or especially North American
counterparts. This is upheld despite the
significant advances made in the recent years
in the research of the European continental
Upper Cretaceous (e.g. Rage, 2002),
especially through continuous fieldwork in
classical areas such as northern Spain or
southern France (e.g. Le Loeuff, 2001), or in
newly discovered localities such as the
Santonian Iharkut locality from Hungary
(Makádi et al., 2006, Ősi & Rabi, 2006). Our
in-depth understanding on the Late Cretaceous European continental ecosystems is
hampered, among other factors, by the almost
complete absence of fortunate taphocoenoses
yielding both plant and animal remains, these
co-occurrences offering rare insights into the
composition and structure of the local
communities (e.g. Calvo et al., 2007) by
preserving mostly autochthonous or parautochthonous taxa derived from the vicinity of
the place of their burial.
In the Maastrichtian of the Haţeg Basin
(Southern Carpathians, Romania) only one
site yielded both plant and animal remains –
the Budurone microvertebrate fossil site (MFS;
see Grigorescu et al., 1999). First recognized
for its relatively abundant and diverse
microvertebrates, the special lithology of the
site also allowed the recovery of a similarly
diverse plant assemblage, with palynomorphs
and
paleocarpological
remains.
The
sedimentology of the site, as well as the
taphonomical features of the local flora and
fauna point to the presence of a
1
taphocoenosis made up of locally derived
components that underwent moderate to very
little transport (except part of the palynoflora).
Consequently, the study of this unique
assemblage allows a thorough reconstruction
of a local paleoenvironmental niche inhabited
by the Haţeg continental biota.
GEOLOGICAL SETTING
The Haţeg Basin is a post-tectonic depression formed in the aftermaths of the Latest
Cretaceous (Second Getic, following Săndulescu, 1984) orogenetic phase of the Alpine
orogenesis, responsible for the thrusting of the
main nappes of the Southern Carpathians. Its
formation is linked to local subsidence
processes occuring around rapidly uplifting
metamorphic core complexes such as the
Retezat Dome south of the basin, the later
becoming place of important molasse-type,
mostly siliciclastic sediment accumulation.
Thick continental sedimentary rock piles
deposited during the Latest Cretaceous are
known in the northwestern and central-eastern
parts of the basin, and were named the
Densuş-Ciula and Sînpetru formations,
respectively (Grigorescu, 1992; Fig. 1).
Although these two units differ somewhat in
lithological composition and overall thickness,
are usually considered as being timecorrelative, a conclusion supported by their
largely comparable paleontological content
(see Grigorescu, 2005, for a recent review).
Based on age determinations of the underlying
marine deposits (Grigorescu & Melinte, 2002,
Melinte & Bojar, 2006, Neagu, 2006),
palynology (Antonescu et al., 1983, van
Itterbeeck et al., 2005) and magnetostratigraphy (Panaiotu & Panaiotu, 2002),
Laboratory of Paleontology, Faculty of Geology and Geophysics, University of Bucharest, 1 N. Bălcescu Blvd., 010041
Bucharest, Romania, dinozoli@geo.edu.ro
2
Geological Institute of Romania, Bucharest, 1 Caransebeş St., 79678, Bucharest, Romania, anapalino@yahoo.com
3
Laboratory of Paleontology, Faculty of Geology and Geophysics, University of Bucharest, 1 N. Bălcescu Blvd., 010041
Bucharest, Romania, dangrig@geo.edu.ro
Z. CSIKI, A. IONESCU & D. GRIGORESCU
the deposits are considered of Maastrichtian
age. The presence of the K/T boundary in the
sedimentary successions was suggested by
Weishampel et al. (1991), but is not yet
supported by conclusive evidence.
The Budurone MFS is located southwest of
Vălioara village (Fig. 1A), one of the most
important fossil localities of the Densuş-Ciula
Formation, being placed within the fossilbearing Middle Member of this unit. The MFS
is located in the upstream half of the Budurone
creek, an ephemeral left-side tributary of the
Vălioara (or Fântânele) creek that flows from
north to south through Vălioara village.
Budurone creek was already known to yield
vertebrate remains as early as 1915 (Kadic,
1916), Kadic excavating reptile remains from
several spots (currently impossible to identify)
located along the creek. More recently,
associated remains of a juvenile individual of
Zalmoxes shqiperorum (Weishampel et al.,
2003) were recovered from Budurone
(Mânzăreanu, 1988, unpubl. B.Sc. thesis),
apparently somewhat downstream, and at a
slightly lower stratigraphic level, than the
Budurone MFS discussed in the present
contribution.
The fossiliferous potential of the Budurone
MFS was first recognized in 1996, being
excavated intermittently beginning with 1998,
through washing and wet-screening of the
fine-grained matrix. Altogether, over 800 kg of
fossiliferous sediment were processed by
standard microvertebrate recovery techniques,
and more material is presently being screenwashed.
The site was mentioned and briefly
described by Grigorescu et al. (1999), who
reported the presence of frog, albanerpetontid,
lizard and crocodylian remains. Besides
vertebrates, shells of continental gastropods
and carpological remains were also recovered
during the screen-washing process, despite
their fragility and the potentially destructive
character of the recovery process. Finally, as
the dark hues of the sediments were
suggestive of an environment with high
potential
for
palynological
material
preservation, it was sampled and yielded a
moderately diverse palynoflora as well.
Lithology and sedimentology of the
Budurone site
The Middle Member of the Densuş-Ciula
Formation is made up of repetitive sequences
of alternating coarse- and fine-grained
siliciclastic deposits, mainly matrix-supported
conglomerates, lithic sandstones and red and
50
grey-green micaceous silts and mudstones;
sorting is generally poor in these deposits, and
coarse-grained terms usually contain high
proportions of angular or slightly rounded
clasts derived from a dominantly metamorphic
source area. Lithoclast petrology, heavy
mineral spectra and sedimentary structures
within the deposits of the Middle Member
suggest a mainly northern source area,
represented by the uplifted metamorphic
basement of the Poiana Ruscă Mts.; minor
contribution of volcanoclastic material is also
noted in several instances, e.g. in the local
section of the Tuştea dinosaur nesting site.
Sedimentological observations suggest that
the deposition of the Middle Member took
place within a dominantly fluvial setting,
represented by alluvial fans as well as
anastomosed and braided rivers.
The Budurone local section (Fig. 1B),
exposed along the bed and steep walls of the
creek, is represented by gray, gray-green or
blackish micaceous silts and silty mudstones,
interbedded with greenish microconglomerates
and coarse sandstones. The coarse-grained
beds have an erosional, irregular base cutting
into the underlying finer sediments; they show
internal size grading and pass either gradually
or sharply into the overlying fine ones. The
thickness of these beds exceeds several
centimetres, ranging up to 50 cm. The finergrained deposits, appearing usually in thicker
beds (up to 1 m), are massive and well
consolidated. Their swelling nature suggests
highly smectitic composition, also supported
by mass spectrometry analyses (work in
progress).
The fossiliferous bed is represented by a
well-sorted, micaceous silty mudstone, over 1
m thick, wedged between two 40-cm thick
lithons of coarse sandstones showing graded
bedding, in the median part of the succession.
The dark-colored, blackish - slightly bluish silty
mudstone is massive, without evident
macroscopic sedimentary structures; it
contains small, framboidal pyrite concretions
and a large quantity of fusinite (charcoal)
fragments, sometimes up to 0.5-0.7 cm long.
The sedimentological interpretation of the
succession suggests the presence of a poorly
drained alluvial plain, with areas of high water
saturation and ponding, but also influenced by
fluvial sedimentary processes from nearby
channels (probably as crevasse splay
deposits, judging from the sheet-like geometry
of the coarser beds and lack of identifiable
channel-linked structures within them).
THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG
BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
Figure 1. A. Simplified geological map of the Haţeg Basin, showing the location of the Budurone MFS near
Vălioara village; inset shows position of the Haţeg Basin. B. Partial lithological log of the Densuş-Ciula
Formation along the Budurone Creek, showing position of the Budurone MFS.
The fossiliferous bed is here interpreted as
a locally developed small pond or marshy
area, based on the structureless massive character of the siltstone and the presence of features suggesting a reducing geochemistry of
the depositional micro-environment (charcoal,
pyrite concretions, plant remains).
FLORAL AND FAUNAL ASSEMBLAGE OF
THE BUDURONE MICROVERTEBRATE
FOSSIL SITE
1. Flora
The plant remains are rare in the
Maastrichtian continental beds of the Haţeg
Basin, especially due to the mostly oxidizing
character of the deposits and to the high
energy of the sediment deposition. Macroplant
remains are especially rare, being confined
mostly to the tuffaceous, fine-grained
lacustrine deposits of the Lower Member of
the Densuş-Ciula Formation (see Grigorescu,
1992), the main localities being reported
around Densuş village (e.g. Mărgărit &
Mărgărit, 1967). Rare isolated plant fragments,
represented by impressions of "Palmophyllumtype" angiosperm leaves, are also known from
the deposits cropping out along the Sibişel
valley at Sînpetru or the Bărbat River valley at
Pui. However, plant remains were represented
up to now only by leaf impressions.
Palynofloras are better known from these
continental deposits (see below); these were
recorded from both the Sînpetru and DensuşCiula formations and were used to constrain
the age of the deposits and to reconstruct the
paleoenvironment the vertebrate fauna lived
in.
1.1. Paleocarpology
The Budurone MFS is unique within the
Maastrichtian of the Haţeg Basin in yielding
carpological remains. These remains are
rather well preserved, despite their fragility,
and the assemblage is relatively diverse.
During the screen-washing of the sediments,
several dozens of seeds and fructifications
were recovered, and were sent for study to E.M. Friis (Stockholm, Sweden) and her coworkers; the peliminary results of their study
shows that the assemblage is interesting.
According to these studies (E.-M. Friis,
written comm., 2000; May, 2003, May et al., in
prep.), the assemblage includes seeds
referable to the angiosperm taxon Eurya sp.,
as well as fruits of Normapolles type
(Normapolles sp.). The largest part of the
assemblage is represented by diverse forms
referred to the carpological metataxon
Carpolithes; about 15 different indeterminate
species of Carpolithes fruit types have been
identified, including nuts, drupes, seeds and
capsules (May, 2003), suggesting a high
diversity of reproductive strategies already
present in the local Maastrichtian flora.
The in-depth study of the Budurone
carpological assemblage is yet to be
undertaken, in order to better understand its
taxic diversity, as well as ecological and
evolutionary significance.
1.2. Palynology
The generally reducing character of the
sediments, along with the presence of carpological remains, charred wood and charcoal
fragments suggested that the Budurone MFS
might have good potential for palynomorph
51
Z. CSIKI, A. IONESCU & D. GRIGORESCU
preservation; the sampling of the fossiliferous
bed yielded indeed a moderately well
preserved palynological assemblage.
Due to their originally large number and
resistance, palynomorphs are easily preserved
in continental settings. In the Maastrichtian
continental deposits of the Haţeg Basin,
although more frequently recovered than
macroplant
remains,
their
restricted
occurrence is still strongly controlled by the
mainly oxidizing character of the sediments.
The first reports on the Latest Cretaceous
palynological record from the Haţeg Basin are
that of Stancu et al. (1980) and Antonescu et
al. (1983). The latter authors identified two
relatively rich assemblages from the DensuşCiula Formation (from the neighborhood of
Vălioara and Densuş, respectively), and a
somewhat less diverse flora from the Sînpetru
Formation at Sînpetru (see also Grigorescu,
1983, 1992). According to this pioneering
study, the assemblage supported a Latest
Cretaceous (Late Maastrichtian) age of the
deposits. It was dominated by fern spores, but
with a diverse component of angiosperm
pollen of Normapolles (and subordinately
Post-Normapolles) type. Another assemblage
was recorded, but not published, from the Pui
area by Ştiucă (1983, unpubl. B. Sc. Thesis).
Subsequently, palynological assemblages
from the Bărbat River (Pui) and Râul Mare
River (Toteşti, Nălaţ-Vad) were studied by van
Itterbeeck et al. (2005). These authors
corrected the previous age assignment based
on palynofloras to Maastrichtian, and noted
the important contribution of the gymnosperm
pollen to the local assemblages, as well as the
dominance of the fern spores and diversity of
the angiosperm pollen.
The samples collected from the Budurone
MFS were treated by standard palynological
techniques. The sample was washed and
dried before mechanical disintegration,
followed by a two-step chemical disintegration
with HCl and HF. The separation of the
organic material was done through heavy
liquid separation using ZnCl2. The resulted
material was mounted on 10 slides and was
studied using an OLYMPUS CX41 transmitted
light microscope (all slides are deposited in the
A. Ionescu collections of the Geological
Institute of Romania).
The recovered palynological assemblage
includes only continental taxa, both spores
and pollen being well preserved. Spores are
subordinate in abundance, while pollen is
dominant both in number and diversity.
Among the spores (Plate I, II), monolete
spores are rare (being represented by
Laevigatosporites ovatus W ILSON & W EBSTER,
52
Cicatricosisporites dorogensis POTONIE &
GELLETICH and Cicatricosisporites sp.), while
trilete spores dominate (about 80% of the
spore assemblage) and include: Appendicisporites tricarinatus, Appendicisporites sp.,
Camarozonosporites insignis NORRIS, Cyathidites australis COUPER, C. sp., Gleicheniidites
sp., Cingulatisporites sp., Leiotriletes sp.,
Lycopodiumsporites clavatoides COUPER, L.
sp., Polypodiaceoisporites sp., Polypodiidites
sp., Rouseisporites simplex ROUSE and
Stereisporites stereoides PFLUG. Fern spores
like Polypodiaceoisporites sp., Polypodiidites
sp. and Cingulatisporites sp. are dominant,
being followed by Gleicheniidites sp.,
Leiotriletes
sp.
and
bryophytes
(Stereisporites).
Both gymnosperm and angiosperm pollen
was recorded. Gymnosperm pollen is rare
(about 5% of the assemblage), with Alisporites
bilateralis ROUSE and Inaperturopollenites dubius PFLUG & THOMSON. Both Normapolles
(Interpollis, Nudopollis, Occulopollis and
Trudopollis) and post-Normapolles (Subtriporopollenites, Triatriopollenites) angiosperm
pollen was recovered. The complete list of the
identified taxa includes: Ephedripites jansonii
(POCOCK) MULLER, Interporopollenites cf.
nagyae KEDVES & HEGEDUS, I. sp., Interpollis
supplingensis (PFLUG) KRUTZSCH, I. sp.,
Labrapollis sp., Minorpollis sp., Nudopollis cf.
terminalis PFLUG & THOMSON, N. thiergartii
(POTONIE) PFLUG, Occulopollis div. sp.,
Pseudovacuopollis
sp.,
Plicapollis
sp.,
Subtriporopollenites annulatus PFLUG &
THOMSON, Triatrioporopollenites sp. and
Trudopollis div. sp. (Plate I, II).
The Budurone local palynoflora differs from
previously known ones from the Haţeg Basin
in the importance (both in diversity and abundance) of the Normapolles and post-Normapolles angiosperm pollen and from those
recorded from the Bărbat and Râul Mare rivers
(van Itterbeeck et al., 2005) by the reduced
participation of the gymnosperm pollen. These
differences probably reflect local environmental
conditions
(see
below,
Paleoenvironment).
2. Fauna
2.1. Invertebrates
Invertebrates are not uncommonly found in
the Maastrichtian deposits of the Haţeg Basin,
but usually these are represented only by
indeterminate paper-thin shell fragments;
screen-washing methodology further contributes to the destruction of the invertebrate
shells. However, in several instances (including the Budurone MFS) a relatively well preserved and diverse invertebrate fauna was
THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG
BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
recovered (Antonescu et al., 1983, Pană et al.,
2002).
At Budurone, invertebrates are represented
exclusively by gastropods (Pană et al., 2002).
The assemblage is dominated by aquatic taxa,
with diverse lymnaeids: Lymnaea maastrichta
PANĂ 2002 (in Pană, Grigorescu, Csiki et
Costea, 2002), L. dilatata PANĂ, 2002, Physa
patula NICOLAS, 1890, Aplexa galloprovincialis
(MATHERON), 1842, and Eustagnicola cretacea
PANĂ, 2002. Besides aquatic taxa, terrestrial
snails are also present, with small sized, but
well-preserved
cyclophorids
[(Rognacia
abreviata (MATHERON), 1832 and Ajkaia cf.
gregari
TAUSCH,
1886)],
vertiginids
(Pyramidula
grigorescui
PANĂ,
2002),
ariophontids (Eoconulus REINHART, 1883) and
helicids (Arionta precursor PANĂ, 2002 - all
new taxa erected in Pană et al., 2002).
The dominance of the aquatic gastropods
suggests the presence of a wet environment,
differing from most other sites yielding
significant
gastropod
faunas
in
the
Maastrichtian of Haţeg (Pană et al., 2002).
Besides gastropods, the Budurone MFS
had yielded a large number of small, ovoidal,
longitudinally ribbed and terminally capped
structures having a shiny black color. These
structures, once thought to be seeds, are
currently interpreted as possible insect eggs
by May (2003). If this inference is confirmed,
then the Budurone site will represent the
second record of Late Cretaceous insects in
the Haţeg paleo-ecosystem (after that
described by Csiki, 2006).
2.2. Vertebrates
Microvertebrates represent an important,
although only poorly known component of the
Late Cretaceous continental vertebrate faunas
of Europe. These were only relatively lately
recognized in the Haţeg Basin (e.g.
Grigorescu et al., 1985), although the recent
years witnessed significant additions to our
knowledge of the local microvertebrate fauna
(see Grigorescu et al., 1999, Codrea et al.,
2002, Smith et al., 2002, Venczel & Csiki,
2003, Folie & Codrea, 2005).
The microvertebrate remains recovered
from the Budurone MFS are not abundant
(slightly over 100 identifiable elements are
known, this number steadily increasing with
continuous work on the site), but interestingly,
they seem to represent a high diversity
assemblage with at least 12 different taxa
already recorded, from fishes to archosaurs.
Fishes
The fish remains are relatively rare in the
Budurone MFS; however, they are better
represented here than in any other
microvertebrate fossil site from the Haţeg
Basin, where these are exceedingly rare
(Fântânele – pers. observ.) or absent
altogether (e.g. Codrea et al., 2002, Smith et
al., 2002). Fishes are represented mainly by
lepisosteids (Plate III, figs. 1, 2); both
Lepisosteus and Atractosteus can be
recognized based on their typical tooth tip
morphology
(pointed
in
Lepisosteus,
lanceolate in Atractosteus; Gayet, in Sigé et
al., 1997). A few rhomboidal, thick, ganoid
scales known from the site can also be
referred to the lepisosteids, a group recorded
from other MFS's in the Haţeg Basin.
However, the presence of another, nonlepisosteid fish can also be suggested, based
on the occurrence of an isolated deeply
amphicoelous fish vertebra, quite unlike the
opisthocoelous vertebrae of the lepisosteids.
This vertebra probably belonged to a (currently
indeterminate) teleostean fish (Peng et al.,
2001).
Anurans
Frog remains are outstandingly abundant
at the Budurone MFS. These are being represented mainly by their characteristic limb elements such as the fused tibiofibulae or radiulnae. Unfortunately, most of the frog remains
are not diagnostic at a low taxonomic level;
consequently, the largest part of the Budurone
frog remains indeterminate (Plate III, fig. 5).
However, Budurone also yielded more
diagnostic elements as well, including the type
specimen of the fossil discoglossid Paralatonia
transylvanica Venczel & Csiki, 2003 (Plate III,
figs. 3-4). It is thus conceivable that at least
part of the indeterminate frog material belongs
also to Paralatonia. However, besides this
discoglossid frog the presence of a second
anuran taxon can also be suggested based on
a fragmentary maxilla, markedly different from
that of Paralatonia.
Albanerpetonids
Albanerpetontids, an extinct group of
allocaudatan amphibians (Fox & Naylor,
1982), are well represented in the Budurone
MFS mainly by their peculiar, characteristic
dentaries with peg-and-socket symphyseal
articulation (Plate III, fig. 6); however, isolated,
fragmentary vertebrae also occur. Fossils
referable to albanerpetontids occur frequently
in the Maastrichtian of the Haţeg Basin (e.g.
Grigorescu et al., 1999, Codrea et al., 2002,
Smith et al., 2002, Folie & Codrea, 2005).
Although formerly (Grigorescu et al., 1999)
some of these remains were referred to the
genus Celtedens, this referral was based on
incomplete elements and is now abandoned.
Consequently, all albanerpetontid remains
53
Z. CSIKI, A. IONESCU & D. GRIGORESCU
from Budurone can be confidently referred to
Albanerpeton (see also Folie & Codrea, 2005).
Turtles
Turtle remains are rare at Budurone,
despite their wide distribution throughout the
Maastrichtian continental beds of the Haţeg
Basin. Turtles are represented only by very
few small, isolated carapace fragments; their
surface ornamentation, made up of fine,
minute, irregular furrows,
is strongly
reminiscent of that of Kallokibotion bajazidi
(Nopcsa, 1923), the only chelonian known
from the Haţeg Basin. Consequently, these
turtle remains are here referred to
Kallokibotion.
Lizards
At Budurone, lizards are uncommon, but
not rare, being represented almost exclusively
by fragmentary jaws. The morphological
diversity of the jaws and dentition suggests the
presence of at least two different lizard taxa.
The first, better documented taxon is
represented by two incomplete maxillae with
widely spaced, slightly tricuspid, cylindrical
teeth with pleurodont implantation (Plate III,
figs. 7-8). The overall morphology of the jaw
fragments suggests scincomorphan affinities.
A fragmentary maxilla preserving a single
almost complete tooth and the largest part of a
second one lacking its tip, represents a
second taxon (Plate III, fig. 9). The robust,
barrel-shaped tooth is somewhat reminiscent
of those of Slavoia described from the Late
Cretaceous of Asia, and is thus referred here
to as ?Slavoia.
Beside these elements, a few isolated
vertebrae and toothless jaw fragments are
also referable to indeterminate lizards
Crocodylians
Crocodylian remains appear to be
exceedingly rare in the Budurone MFS, being
represented only by two isolated teeth; this
contrasts
markedly with the widespread
presence of the crocodylian teeth in the
Maastrichtian deposits of Haţeg, either in
macrovertebrate fossil sites, MFS's or as
isolated finds.
One of these teeth is highly conical, almost
straight and pointed, with small and smooth
mesial and distal carinae (Plate III, fig. 10). It is
reminiscent of the teeth generally assigned to
the eusuchian Allodaposuchus precedens
(Buscalioni et al., 2001; Delfino et al., 2008)
and is accordingly referred to this taxon.
The other tooth is short, widely triangular
and labio-lingually compressed (Plate III, fig.
11). It is similar to teeth referred previously by
Grigorescu et al. (1999) to as Doratodon sp.
and most probably represents a second taxon
54
of crocodylians in the Budurone local
assemblage.
Theropods
Theropods are extremely rare in the
Budurone MFS, being represented by only one
isolated tooth crown (Plate III, fig. 12). Its
morphology is similar to that of the tooth genus
Euronychodon (Antunes & Sigogneau-Russell,
1991), previously reported from other MFS's in
the Haţeg Basin (Csiki & Grigorescu, 1998,
Codrea et al., 2002), with labially convex and
lingually flat sides. Both the mesial and the
distal carinae are smooth and placed on the
lingual face of the crown.
Ornithopods
As in the case of the theropods, ornithopod
remains are rare in the Budurone MFS (only
two isolated shed tooth crowns), although they
occur frequently in other MFS's, as well as in
macrovertebrate fossil sites or as isolated
finds. Despite this low frequency, both ornithopod taxa known from the Haţeg Basin (Telmatosaurus, Zalmoxes - Plate III, fig. 13;
Grigorescu, 2005) are represented, based on
their very characteristic tooth morphologies
(Weishampel et al., 1993, 2003).
Eggshells
Eggshell fragments, not exceeding 3-4 mm
in maximum dimension, are only uncommonly
found in the Budurone MFS. Several
morphotypes can be identified. One of these
represents the megaloolithid morphotype,
relatively thick eggshell fragments with the
outer surface covered by large tubercles, also
known in the dinosaur eggs reported from
several sites spread around the Haţeg Basin
(e.g. Grigorescu et al., 1994, Codrea et al.,
2002, Smith et al., 2002). Eggs with this shell
morphotype are usually linked to titanosaurian
sauropods (e.g. Chiappe et al. 1998), but are
considered to belong to the hadrosaur
Telmatosaurus in the Haţeg Basin. Another
eggshell morphotype is thinner (not more than
1 mm in thickness) and with the outer surface
covered with sparse, crater-like tubercles,
being described from the Upper Cretaceous of
southern France as geckonoid eggshells
(Garcia, 2000). A third morphotype is
represented by thin eggshells, delicately
ornamented with small tubercles; these might
represent theropod eggshells.
TAPHONOMICAL CHARACTERISATION OF
THE BUDURONE MFS
The taphonomy of the Maastrichtian
continental beds from Haţeg is studied only in
a preliminary manner, the most important
contributions concerning the characteristic
macrovertebrate fossil sites, the so-called
"fossil pockets" (see Grigorescu, 1983).
THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG
BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
Recently, a thorough study of the taphonomy
of several important fossil accumulations was
undertaken by Csiki (2005, unpubl. Ph.D.
thesis), and the conclusions of this study will
be published elsewhere; thus, the detailed
taphonomical study of the Budurone MFS is
outside of the scope of this contribution.
The most important taphonomical features
will be, however, discussed shortly, in order to
gain insights into the nature of the original
depositional environment and the potential
taphonomical biases that operated.
The taphonomical study of the MFS's
(including Budurone) requires a specific
approach, due to the recovery method of the
fossils, through bulk sediment sampling and
screen-washing. The recovery procedure
implies first of all the almost complete loss of
information concerning the spatial distribution,
orientation or possible association/articulation
of the different specimens. Moreover, the recovery procedure is destructive, leading to
further fragmentation of the more fragile specimens, and partial destruction of the original
taphonomic modifications. Thus, the taphonomic information gained from MFS's is biased
towards the compositional features (abundance, diversity) and against the detailed
understanding of the taphonomic processes
operating.
Taking into account this bias, a good
amount of taphonomic information is still
recoverable, allowing the reconstruction of the
major aspects of the local assemblage's
taphonomic history.
Taphonomic
characteristics
and
modifications
Several hundred microvertebrate remains
were recovered; however, the largest part of
these are indeterminate bone fragments, while
only slightly over 100 specimens are
determinable at least at major clade (e.g.
anuran, lizard) level. The microvertebrate
remains and associated flora and fauna seem
to be concentrated into the upper half of the
fossiliferous bed.
The bone assemblage shows a low
dimensional variability (many specimens have
maximum dimensions of less than 1 mm, and
only a minor percent reaches a maximum
diameter of over 5 mm); thus, size sorting
seems to be good to excellent. Morphological
diversity is, by the contrary, rather high;
although long tubular bones dominate (somewhat more than 44%), isometric (cube-shaped
– 19,6%) and irregular (15,6%) ones are also
common. The unusual rarity of conical elements is due to the scarcity of isolated teeth.
Hydraulic compatibility between the finegrained sediments and the dominantly small-
sized microvertebrate remains suggests that
the same agents are responsible for the
accumulation of both the sediments and the
fossils; the same compatibility is obvious in the
case of the carpological and invertebrate
remains as well.
Taphonomical modifications are represented mainly by fractures; pre-burial and
diagenetic fractures as well as fractures
related to the recovery process are all present
and sometimes difficult to separate. Bone
splinters are frequent. Weathering and
abrasion of the microvertebrate remains is
variable;
while the better preserved,
taxonomically determinate specimens are only
slightly weathered and abraded, if at all, the
indeterminate bone chips and splinters are
usually heavily weathered so as to expose the
internal bone structure and are also highly
abraded. The weathering and abrasion stages
shown by the bone fragments suggest these
experienced repeated episodes of transport
and reworking. This was not, however, the
case of the better-preserved remains. Besides
being significantly less abraded and
weathered (showing the unaltered smooth
shiny outer cortex of the bones), these also
include extremely fragile elements such as
tooth-bearing jaws fragments of amphibians or
lizards. The above observations suggest that
this component of the thanatocoenosis
experienced either only very short transport
and short-term subaerial exposure or none at
all (excepting possibly the rare isolated teeth
that, being among the most resistant elements
of the skeleton, can better withstand transport
and subaerial exposure).
Composition, abundance and diversity
Although based only a limited sample
(slightly more than 100 specimens), the
vertebrate assemblage preserved in the
Budurone MFS is relatively diverse, as it
includes at least 12 different vertebrate taxa.
The most diverse group is that of the fishes,
with 3 different taxa, which is outstanding
compared to other Haţeg local assemblages in
which fish remains are extremely rare or
absent altogether. Ornithopods, crocodylians,
anurans and lizards are slightly less diverse
(each being represented by 2 taxa – maybe 3,
in the case of the frogs), while theropods,
turtles and albanerpetontids are the least
diverse clades, with only one taxon present.
The taxic abundance is estimated based on
the minimum number of identified specimens,
as proposed previously for microvertebrate
sites consisting mainly of disarticulated and
isolated remains (e.g. Foster, 2001, 2003); this
approach assumes that each element
represents another individual, as skeletal
55
Z. CSIKI, A. IONESCU & D. GRIGORESCU
association cannot be proved. Based on this
count, anurans are the most abundant group
(about 75% of the identifiable remains),
followed by the lizards (10%), albanerpetontids
(5%) and fishes (4%). On the other end of the
abundance spectrum, the crocodylians,
theropods, ornithopods and turtles are rare
(under 2% NIS).
However, confronting the abundance and
diversity data shows some interesting
patterns. While the material referable to the
two (or possibly three) anuran taxa totals
about three-quarters of the NIS, the fishes,
with a comparable taxic diversity figure,
yielded only 4 specimens (less than 4%).
Similarly, the taxonomically least diverse
albanerpetontids are represented by several
specimens (5%), but the relatively more
diverse crocodylians or ornithopods are
identified based on very few remains (less
than 2%).
All major skeletal element categories (skull
fragments, isolated teeth, vertebrae, girdle,
limb and dermal elements) are represented,
although the elements of the appendicular
skeleton dominate by far (over 44%); this is
mainly due to the over-representation of the
anuran tibiofibulae, humeri and phalanges.
Jaw fragmens and vertebrae are also relatively
common (16,7%), while other cranial elements
(1%) and girdles (2%) are rare. Especially
noteworthy is the extreme rarity of the isolated
teeth (of dinosaurs and crocodylians) and
dermal elements (reduced to turtle shell
fragments); both of these categories occur
commonly in other MFS's.
Distribution of the different skeletal categories among the major taxonomic groups
represented shows a pattern that occurs
frequently in other MFS's in the Haţeg area:
typical microvertebrates such as anurans, and
to a lesser extant albanerpetontids and lizards
(as well as the otherwise almost unknown
fishes) are represented by a wide range of
skeletal elements from fragile cranial to more
robust vertebral elements, while macrovertebrates are known almost exclusively by a
single skeletal category: isolated teeth
(crocodylians, dinosaurs) or dermal plates
(turtles). In this last case, the categories
involved include the most resistant elements of
the skeleton, suggesting that macrovertebrate
remains underwent specific taphonomic
processes selecting (besides a normal sizerelated bias) for those remains that best
withstand weathering, breakage and abrasion.
Discussions
The Budurone MFS is among the most
important ones in the Haţeg Basin, mainly not
because of the quantity of the recovered
56
microvertebrate remains, but of its special
taphonomic signature.
The
most
remarkable
taphonomic
characteristics of the Budurone MFS are
represented by the: a) quasi-absence of
subaerial weathering and abrasion in the
taxonomically identifiable elements; b) good
preservation state of fragile skeletal elements
such
as
toothed
jaw
fragments,
albanerpetontid vertebrae, but also the
angiosperm seeds and fruits or the relatively
well preserved thin-shelled gastropods etc.; c)
low frequency of the otherwise resistant
skeletal elements, commonly recovered from
other MFS's from Haţeg, such as isolated
archosaur teeth or turtle plates; d) high
frequency of indeterminable bone fragments
and scraps showing complex fracturing, as
well as advanced weathering and abrasion.
Together, these features suggest that the
Budurone assemblage represents a composite
attritional taphocoenosis, including two
different parts – one with a high degree of
taphonomical elaboration (represented by the
bone fragments) and one with low degree of
taphonomical alteration (measured either by
weathering, abrasion or fracturing). This
second component most probably represents
a
locally
derived,
autochthonous
or
parautochthonous assemblage, while the
taphonomically
altered
component
is
allochthonous, and probably reworked in
several phases. Consequently, the wellpreserved component of the taphocoenosis
(and the one that includes the largest part of
the taxonomically identifiable elements)
reflects a local biocoenosis, with a limited
amount of spatial and time averaging.
The assemblage is politaxic with a large
number of individuals represented, and
taxonomically relatively diverse. However, it
shows some interesting pattern such as:
a) low intra-clade diversity (between 1 and
maximum 3 taxa in each group, including
otherwise diverse clades such as lizards,
theropods or crocodilians);
b) several compositional peculiarities,
compared to other Haţeg MFS's, such as the
extreme abundance of the frog remains,
relative high diversity of the fishes (otherwise
almost unknown from other MFS's), as well as
the rarity of turtle, crocodylian and dinosaur
remains (usually more frequently found in the
different MFS's)
c) a distinction between two different
categories of the identifiable remains, based
on skeletal representation. The first category
(A), illustrated first of all by frogs, but to a
lesser extent also by albanerpetontids, fishes
and lizards, is represented by taxa with
THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG
BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
numerous and diverse skeletal elements
preserved, belonging to different parts of the
body and including extremely fragile skeletal
parts in good preservation state. The second
category (B) includes the turtles and
archosaurs, represented by few elements from
only one skeletal category, usually one with
high preservation potential.
d) the differential preservation is also replicated by differences in size (category A
includes small-sized taxa, while category B
larger-sized ones) and ecological preferences
(category A are mainly strictly aquatic taxa,
while category B mainly strictly terrestrial, or
semiaquatic, but not water-bound ones).
This twofold division of the identifiable
remains in the Budurone assemblage
suggests that it is formed by two distinct
biocoenotic components. One is locally
derived, autochthonous, with no or extremely
reduced transport from the microhabitat it
actually inhabited, the skeletal remains being
buried in large numbers shortly after the death
of the organisms, within their living
environment. This component also includes,
besides most of the category A vertebrate
remains (excepting possibly the lizards), also
the carpological remains and gastropods as
well.
The
other
component
is
a
parautochthonous one, derived from nearby
microhabitats, but without much reworking or
long-distance transport; occasional sediment
influx is probably responsible for the
introduction of some of these components, as
suggested by the hydraulic compatibility
between
them
and
the
surrounding
sedimentary matrix. However, selection of the
remains occurred during this transport by
preferential concentration of the more resistant
elements such as isolated teeth or shell
fragments (category B elements). One
exception to this is represented by the lizard
remains, included in category A, but derived
from strictly terrestrial taxa, suggesting the
lizards inhabited closely located microhabitats
and underwent extremely short transport,
while other terrestrial taxa lived farther from
the site of the burial. The semi-aquatic turtles
and crocodilians, rare in the Budurone MFS,
despite their common occurrence in other
MFS’s, represent another partial exception.
This is probably explained by the fact that the
specific depositional environment at Budurone,
although wet, was not particularly suitable for
these taxa.
DISCUSSIONS
Faunal composition of the Budurone MFS
The composition of the Haţeg paleobiocoenosis is now relatively well-known (see
Grigorescu, 2005), although discovery of new
taxa (especially invertebrates, plants, but also
vertebrates) is to be expected. Different sites
yielded local assemblages with slightly
different compositions, but usually the nature
of the main groups represented and their
relative participation are largely predictable.
The Budurone MFS, however, yielded a rather
peculiar floral and faunal assemblage, with
taxonomic composition and especially relative
abundance of the taxa that set it apart from
other sites of the Haţeg area.
Fishes are extremely rare in the Haţeg
fauna, besides Budurone being reported from
only two other MFS's: from the Pui locality
excavated by a Romanian-French team
(Grigorescu et al., 1985) and from FântâneleVălioara (Grigorescu et al., 1999). At Pui,
acipenseriform and characid fish remains were
found, while Fântânele yielded lepisosteid and
possible characid remains. The rarity of fish
remains in Haţeg contrasts markedly with their
common occurrence in other European Late
Cretaceous faunas where lepisosteids are
common, and usually extremely abundant,
while other taxa are also present (e.g. Cavin,
1999, Makádi et al., 2006). In this respect, the
Budurone MFS is outstanding in showing the
highest fish diversity with three different taxa,
although their remains are rare.
Anuran remains occur frequently in the
different MFS's around the Haţeg Basin (e.g.
Grigorescu et al., 1999, Codrea et al., 2002,
Smith et al, 2002, Folie & Codrea, 2005).
Although usually not identifiable to lower
taxonomic level, their very characteristic
remains are easily recognizable and thus allow
the identification of the clade. Most anuran
remains are referred to different taxa within the
Discoglossidae (Grigorescu et al., 1999,
Venczel & Csiki, 2003, Folie & Codrea, 2005),
but other taxa were also reported (Venczel &
Csiki, 2003). At Budurone, several anuran
remains are referable to the Discoglossidae,
and some of them to the discoglossid
Paralatonia transylvanica (Venczel & Csiki,
2003), the type locality of which is represented
by Budurone. Paralatonia is also known from
other MFS's from the Haţeg Basin. The taxon
is not yet reported from outside the Haţeg
area. Besides Paralatonia, another anuran
taxon is also present; it is rare at Budurone,
and not yet discovered in other MFS's.
Fossils referable to the extinct group of
allocaudatan albanerpetontids occur frequently
in the Maastrichtian of the Haţeg Basin (e.g.
Grigorescu et al., 1999, Codrea et al., 2002,
Smith et al., 2002, Folie & Codrea, 2005).
Although formerly (Grigorescu et al., 1999)
some of these remains were referred to the
57
Z. CSIKI, A. IONESCU & D. GRIGORESCU
genus Celtedens (now known to be restricted
to the Early Cretaceous, McGowan, 2002), this
referral was based on incomplete elements
and are now abandoned. Consequently, all
albanerpetontid remains can now be
confidently referred to Albanerpeton (see also
Folie & Codrea, 2005), a genus with a wide
stratigraphic distribution, ranging from the
Aptian to the Pliocene (Gardner, 2002,
Gardner et al., 2003, Venczel & Gardner,
2005). Albanerpetontids, mostly indeterminate
taxonomically, are reported from the
Santonian of Hungary (Makádi et al., 2006) as
well as from several western European Late
Senonian localities (e.g. Buffetaut et al., 1999,
Duffaud & Rage, 1999, Garcia et al., 2000).
The turtle Kallokibotion is frequently
occurring in the fossiliferous sites in the Haţeg
Basin, and it also present in the Budurone
MFS. However, its remains are unusually rare
here, while they are common in most MFS's,
as well as macrovertebrate sites. Kallokibotion
is not yet reliably reported from the Late
Cretaceous of Europe outside Romania
(Gaffney & Meylan, 1992, Codrea & Dica,
2005), except maybe from the Lower Campanian of Austria (Sachs & Hornung, 2006).
Lizards are a recently discovered
component of the Haţeg local fauna, being first
reported by Grigorescu et al. (1985), but
described on some detail only by Grigorescu
et al. (1999). Since then, several important
contributions were published (Codrea et al.,
2002, Smith et al., 2002, Folie & Codrea,
2005) revealing a diverse lizard component of
the assemblage. Lizard remains are commonly
occurring in different microvertebrate fossil
sites (Grigorescu et al., 1999, Codrea et al.,
2002, Smith et al., 2002, Folie & Codrea,
2005), but few taxa were discovered in more
than one site. The local diversity of the lizards
rarely exceeds two to four taxa, this being the
case also at Budurone. One of the taxa
represented is unknown from other MFS's,
while the other one (?Slavoia) is probably
present at Pui as well. None of these taxa
were previously reported from other Late
Cretaceous European localities, although our
knowledge both about the lizards from Haţeg
and Europe is still admittedly poor.
Both Allodaposuchus and Doratodon occur
frequently in different MFS's in Haţeg.
Allodaposuchus is rather common also in
macrovertebrate sites or as isolated finds,
while Doratodon, probably a smaller-sized
taxon based on its known remains, is more
rarely found in these circumstances. Their
remains are, however, surprisingly rare in the
Budurone MFS. Allodaposuchus is a wideranging
taxon,
known
from
other
58
Transylvanian sites outside Haţeg (Codrea &
Dica, 2005; Delfino et al., 2008), but also from
southern France and Spain (Buscalioni et al.,
2001). Doratodon is also known outside
Romania, from the Santonian of Hungary (Ősi
& Rabi, 2006), as well as the Campanian of
Austria (Seeley, 1881) and Spain (Company et
al., 2005).
The rarity of dinosaur remains in the
Budurone assemblage, reduced to but one
isolated tooth for each taxon represented, is
again remarkable, compared to other MFS's
from the Haţeg Basin. Euronychodon is a
small theropod, known only by its distinctive
teeth, being recovered from several sites in
Haţeg (Csiki & Grigorescu, 1998, Codrea et
al., 2002). It is also known from several
western European localities, both from the
Lower (Rauhut, 2002) and the Upper
Cretaceous (Antunes & Sigogneau-Russell,
1991) of Iberia. However, the absence of
dromaeosaurid teeth, common in other sites,
is surprising.
On the other hand both Zalmoxes and
Telmatosaurus, the ornithischians usually
represented in other fossiliferous sites from
the Haţeg Basin, including MFS's as well, are
present at Budurone. They are among the
most common dinosaurian taxa from the
Haţeg Basin and surrounding areas (pers.
obs.; Codrea & Dica, 2005). Zalmoxes is
probably endemic to the Transylvanian area,
but is possibly also present in the Lower
Campanian of Austria (Sachs & Hornung,
2006); Telmatosaurus, however, seems to be
restricted to the Transylvanian area and
unknown from more western parts of Europe
(Laurent et al., 1997).
In overall, the composition of the Budurone
local assemblage is comparable to that of
other Haţeg MFS's, but differs from all of these
in the presence of a diverse fish component, in
the rarity of the terrestrial archosaurs and
overwhelming abundance of the frogs.
Moreover, it is as yet unmatched in that
invertebrates and especially plant remains
(palynomorphs, fruits and seeds) are
associated to the microvertebrates. This
unique composition is probably due to the
unique microhabitat represented by the
depositional environment of the fossiliferous
bed.
Biostratigraphy
From the Budurone local assemblage, the
palynomorphs are especially useful as age
indicators. Most of the recorded palynomorph
taxa have a Late Cretaceous-Early Tertiay
range, but typical Paleocene taxa are missing.
The dominance and diversity of the
Normapolles group, especially the presence of
THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG
BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
such
taxa
as
Interpollis,
Nudopollis,
Occulopollis and Trudopollis, suggest a
Maastrichtian age for these deposits (Goczan
et al., 1967, Batten, 1981, Portiagina, 1981,
Herngreen et al., 1986), similar to those
studied by van Itterbeeck et al. (2005).
Paleoenvironment
Several lines of evidence point to the presence of a particular depositional environment
at Budurone, one that differs from those
represented by other MFS's. These features
include: the dominance of aquatic gastropods,
the preservation and high diversity of
angiosperm seeds and fruits (duplicated by the
high diversity and dominance of angiosperm
pollen), the dominance of the remains of
aquatic-semiaquatic taxa (especially frogs),
the unusual diversity of fishes, the rarity of
typical terrestrial taxa, represented exclusively
by remains with high preservation potential
and the predominantly local derivation
(autochthony)
of
the
microvertebrate
thanatocoenosis, as suggested by the
taphonomic features. The sedimentological
character complex, especially the structureless massive dark gray fine-grained sediments
with high frequency of charcoal fragments and
pyrite concretions, is also unknown from other
sites.
Together, the sedimentological, paleontological and taphonomical evidence points the
presence of a wet, water-logged or ponded
environment, probably a small swamp or
shallow, marshy pond microhabitat within a
poorly drained floodplain.
The lack of any dinoflagellate cysts
suggests a typical continental setting for the
Budurone depositional site, weakening the
proposed arguments for the presence of
marine or brackish conditions in the Vălioara
area (e.g. Vremir, 1998).
The palynological assemblage points to the
presence of a warm, tropical-subtropical
climate, suggested by the rarity of bisaccate (3
Alisporites specimens) and taxodiacean (one
Inaperturopollenites species) pollen, the common presence of ferns such as Gleicheniidites,
Lygodiumsporites, Cicatricosisporites and
Cyathidites. The presence of a somewhat drier
subtropical climate is also supported by the
abundance of the Normapolles pollen group
(Pacltova, 1966, Zaklinskaya, 1977).
The high diversity of Normapolles pollen,
but especially that of the angiosperm seeds
and fruits (whose good preservation, despite
their fragility, suggests short transport and
thus autochthony) allows the reconstruction of
the local vegetation as being closed, forested,
made of a thick angiosperm ground cover and
low canopy, with participation of herbaceous
and tree ferns, a vegetation different both from
the mangrove vegetation reconstructed for the
nearby Rusca Montană Basin (Petrescu &
Duşa, 1980) or the more open, wooded
savannah vegetation of the Râul Mare-Râul
Bărbat area (van Itterbeeck et al., 2005).
Paleoecology
Taphonomical observations on the Budurone microvertebrate assemblage suggest the
mixture of two different paleocommunities –
one autochthonous to the depositional
environment and one parautochthonous,
derived
from
nearby
areas.
The
autochthonous paleobiocenosis is represented
mainly by aquatic and semi-aquatic taxa, such
as fishes, frogs and albanerpetontids,
inhabiting the pond or swamp and its thickly
vegetated shore forests. Based on the iliac
morphology, Paralatonia was considered to be
a frog with good swimming and/or jumping
abilities, inhabiting riverine or marsh
environments (Venczel & Csiki, 2003).
Albanerpetontids, more terrestrial in habitat,
were showed to live in densely forested areas
as well (Venczel & Gardner, 2005). Lizards,
although terrestrial in habitat preference, might
have also formed elements of this microhabitat, populating the shore forests, based on
the presence of their fragile, but well-preserved toothed jaw fragments. The other taxa,
such as the terrestrial dinosaurs, were inhabitants of nearby drier, more terrestrial habitats,
but probably ventured into the forested shores
of the pond either for browsing or hunting.
The rarity of crocodylian and turtle remains
– both semi-aquatic – is somehow puzzling,
and probably means that these taxa were not
perfectly adapted to the local conditions and
favored more open, lacustrine or fluvial
settings. The absence of the multituberculates,
occurring in many other MFS's, is also
unexpected, especially since the high diversity
of angiosperms would have represented a rich
food source to these taxa. The small number
of identifiable remains might account for their
apparent absence, or, alternatively, they as
well would have preferred drier habitats.
The image of a complex local food web is
emerging from the study of the Budurone
MFS. The base of the web is represented by a
rich angiosperm assemblage, completed by
water-loving ferns and bryophytes. The
vegetation represented the food source for
invertebrates and probably for herbivorous
dinosaurs entering this pond-shore forest from
their preferred drier habitats nearby. The
preserved
autochthonous
invertebrates,
gastropods and possibly insects, were forming
a link between the primary producers and
secondary invertivorous consumers such as
59
Z. CSIKI, A. IONESCU & D. GRIGORESCU
lizards, frogs, albanerpetontids and possibly
some fishes. The top of the local food web
was represented by predatory fishes such as
lepisosteids,
along
with
rare
and
parautochthonous
crocodylians
and
theropods, visiting intermittently this highproductivity and biomass-rich area.
CONCLUSIONS
The discovery of the Budurone MFS adds
important information to our knowledge about
the Maastrichtian continental ecosystem of the
Haţeg Basin. It represents a particular depositional environment, not yet documented by
other MFS's: a wet, heavily forested marshland or swampy pond preserving a good
sample of the local and more distal
paleobiocoenoses. The local paleobiocoenosis
included a rich angiosperm assemblage,
documented by pollen, but also by seeds and
fruits, extremely rarely preserved in the Late
Cretaceous fossil record and otherwise as yet
unknown in the Haţeg area. This type of floral
assemblage differs from those previously
reconstructed, suggesting the presence of a
complex, mosaic-like vegetation pattern. The
Budurone local assemblage also supports
previous paleoclimatic and biostratigraphic
interpretations. The microvertebrates, together
with the invertebrates and plants, allow the
reconstruction of a complex local food web,
dominated by small-sized aquatic and semiaquatic invertivores.
ACKNOWLEDGMENTS
The authors wish to thank all the
participants of the Haţeg field campaigns who
contributed to the excavation of the Budurone
MFS and sorting of the specimens. Dr. M.
Venczel helped in the identification of the
anurans, albanerpetontids and lizards; his
drawings are reproduced in Plate III. The SEM
photographs were taken in Bucharest (by Dr.
C. Costea) and Bruxelles (Julien Cillis). The
suggestions of the anonymous reviewers are
also thanked. The fieldwork was supported by
the CNCSIS grants G239/1998-2001 and
G1163/2004-2006, as well as EC FP5 Project
ABC 95/2003.
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THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG
BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT
PLATE CAPTIONS
PLATE I
Fig. 1. Appendicisporites sp.: slide 4; 10.9/145.2; 55 .
Figs. 2-3. Cyathidites australis Couper: fig. 2 - slide 1; 22.2/133.9; 45.0 ; fig. 3 - slide 3; 12.0/134.0; 27.5 ;
Fig. 4. Rouseisporites simplex Cookson & Dettmann: slide 1; 20.0/138.0 ; 60.0  ;
Figs. 5-11. Polypodiaceoisporites sp. and Cingulatisporites sp.: slide 3; 5; 32.5-45.0 ;
Figs. 12-15. Stereisporites steroides Pflug: slide 2-4; 20-32.5 ;
Figs. 16-18. Gleicheniidites sp.: slide 4; 37.5-42.5 ;
Fig. 19. Laevigatosporites ovatus Wilson & Webster: slide 5; 18.2/138.0; 40 ;
Fig. 20. Cicatricosisporites dorogensis Potonie & Gelletich: slide 4; 6.2/136.0; 52.5 ;
Figs. 21-22. Lycopodiumsporites clavatoides Couper: fig. 21 - slide 5; 8.0-140.1; 47.5 ; fig. 22 - slide 3; 16.2143.0; 25 ;
Figs. 23-24. Taxon indeterminate: fig. 23 - slide 8; 14.1/133.0; 37.5 ; fig. 24 - slide 2; 20.0/134.9; 25.0 ;
Figs. 25-31. Trudopollis sp.: fig. 25 - slide 1; 22.9/141.0; 25.0 ; fig. 26 - slide 5; 10.2/133.0; 17.5 ; fig. 27 slide 8; 15.8/136.9; 20.0 ; fig. 28 - slide 9; 18.0/135.0; 42.5 ; fig. 29 - slide 5; 10.0/149.0; 17.5 ; fig. 30 slide 5; 10.0/138.0; 17.5 ; fig. 31 - slide 1; 24.9/140.2; 17.5 ;
Fig. 32. Taxon indeterminate: slide 5; 6.0/135.0; 42.5 ;
PLATE II
Figs. 1-2. Trudopollis sp.: fig. 1 - slide 5; 13.0/138.9; 15.0 ; fig. 2 - slide 9; 16.1/141.9; 17.5 ;
Fig. 3. Interpollis supplingensis (Pflug) Krutzsch: slide 10; 18.9/149.1; 22.5 ;
Fig. 4. Interpollis sp.: slide10; 15.6/138.0; 12.5 ;
Figs. 5-7. Plicapollis sp.: fig. 5 - slide 1; 23.9/147.0; 20.0 ; fig. 6 - slide 1; 23.9/138.0; 22.5 ; fig. 7 - slide 10;
3.9/146.2; 20.0 ;
Fig. 8. Taxon indet.: slide 5; 6.9/128.9; 32.5 ;
Fig. 9. Trudopollis sp.: slide 10; 19.2/143.1; 15.0 ;
Fig. 10. Minorpollis sp.: slide 9; 13.9/138.9; 15.0 ;
Fig. 11. Polypodiisporites sp.: slide 5; 5.1/145.1; 40.0 ;
Fig. 12. Taxon indet.: slide 5; 6.1/135.2; 45.0 ;
Fig. 13. Subtriporopollenites anulatus Pflug & Thomson: slide 6; 20.1/140.9; 32.5 ;
Fig. 14. Cicatricosisporites sp.: slide 4; 6.2/136.0; 37.5 ;
Fig. 15. Leiotriletes sp.: slide 8; 15.0/130.0; 32.5 ;
Fig. 16. Taxon indeterminate: slide 5; 7.8/135.0; 37.5 ;
Figs. 17-18. Alisporites bilateralis Rouse: fig. 17 - slide 3; 8.2/132.0; 22.5; fig. 18 - slide 3; 16.2/130.5; 42.5;
Fig. 19. Inaperturopollenites dubius Pflug & Thomson: slide 3; 21.0/147.0; 50.0;
Fig. 20. Ephedripites jansonii (Pocock) Muller: slide 3; 11.2/139.4; 80.0;
Figs. 21, 25. Occulopollis sp.: fig. 21 - slide 9; 10.9/138.0; 17.5; fig. 25 - slide 3; 13.0/139.1; 17.5;
Figs. 22-24. Nudopollis thiergartii Pflug: fig. 22 - slide 5 10.5/144.0;20.0; figs. 23-24 - slide 3; 32.0/134.0;
20.0
Figs. 26-28. Occulopollis sp.: fig. 26 - slide 5; 9.9/129.0; 20.0; fig. 27 - slide 2; 10.0/138.0; 20.0; fig. 28 slide 10; 12.8/143.0; 17.5;
Figs. 29-31. Trudopollis sp.: fig. 29 - slide 3; 13.1/139.1; 17.5; fig. 30 - slide 3; 9.1/139.0; 15.0; fig. 31 - slide
4 9.9/142.0; 45;
Fig. 32. Plicapollis sp.: slide 10; 17.9/138.1; 17.5.
PLATE III
Figs. 1-2. Lepisosteidae: fig. 1 – FGGUB v.401, scale, external view; fig. 2 – Lepisosteus sp., FGGUB v.400,
tooth
Figs. 3-4. Paralatonia transylvanica Venczel et Csiki, 2003: fig. 3 – FGGUB v.452, fragmentary ilium, right lateral
view (from Venczel & Csiki, 2003); fig. 4 – FGGUB v.455, holotype, fragmentary ilium, left lateral view
Fig. 5. Anura indet.: FGGUB v.438, sacrum, dorsal view
Fig. 6. Albanerpeton sp.: FGGUB v.414, symphyseal fragment of right dentary, dorsal and lingual views.
Figs. 7-8. Scincomoprha indet.: fig. 7 - FGGUB v.481, fragmentary maxilla, labial view; fig. 8 - FGGUB v.482,
fragmentary maxilla, lingual view.
Fig. 9. Slavoia sp. (?): FGGUB v.470, fragmentary maxilla, lingual view
Fig. 10. Allodaposuchus precedens Nopcsa, 1929: FGGUB unnumbered, isolated tooth, labial view
Fig. 11. Doratodon sp.: FGGUB unnumbered, isolated tooth, lingual view
Fig. 12. Euronychodon sp.: FGGUB R.1826, isolated tooth, lingual view
Fig. 13. Zalmoxes sp.: FGGUB R.1812, isolated dentary tooth, lingual view.
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PLATE I
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PLATE II
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PLATE III