the budurone microvertebrate site from the maastrichtian of the

ACTA PALAEONTOLOGICA ROMANIAE V. 6 (2008), P. 49-66.

1 Laboratory of Paleontology, Faculty of Geology and Geophysics, University of Bucharest, 1 N. Bălcescu Blvd., 010041 Bucharest, Romania, [email protected] 2 Geological Institute of Romania, Bucharest, 1 Caransebeş St., 79678, Bucharest, Romania, [email protected] 3 Laboratory of Paleontology, Faculty of Geology and Geophysics, University of Bucharest, 1 N. Bălcescu Blvd., 010041 Bucharest, Romania, [email protected]

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 Creta-ceous 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 par-autochthonous 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

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 dep-ression formed in the aftermaths of the Latest Cretaceous (Second Getic, following Săndu-lescu, 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 time-correlative, 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 magneto-stratigraphy (Panaiotu & Panaiotu, 2002),

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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 fossil-bearing 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 screen-washed.

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

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 finer-grained 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).


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The fossiliferous bed is here interpreted as a locally developed small pond or marshy area, based on the structureless massive cha-racter of the siltstone and the presence of fea-tures 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 "Palmophyllum-type" 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 Densuş-Ciula 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 co-workers; 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 carpo-logical remains, charred wood and charcoal fragments suggested that the Budurone MFS might have good potential for palynomorph

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.


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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 Densuş-Ciula 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 WILSON & WEBSTER,

Cicatricosisporites dorogensis POTONIE & GELLETICH and Cicatricosisporites sp.), while trilete spores dominate (about 80% of the spore assemblage) and include: Appendici-sporites tricarinatus, Appendicisporites sp., Camarozonosporites insignis NORRIS, Cyathi-dites 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 du-bius PFLUG & THOMSON. Both Normapolles (Interpollis, Nudopollis, Occulopollis and Trudopollis) and post-Normapolles (Subtripo-ropollenites, 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 abun-dance) of the Normapolles and post-Norma-polles 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 environ-mental 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 contri-butes to the destruction of the invertebrate shells. However, in several instances (inclu-ding the Budurone MFS) a relatively well pre-served and diverse invertebrate fauna was


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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, non-lepisosteid 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 repre-sented mainly by their characteristic limb ele-ments such as the fused tibiofibulae or radi-ulnae. 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


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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

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 ornitho-pod taxa known from the Haţeg Basin (Tel-matosaurus, 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).


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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 reco-very procedure is destructive, leading to further fragmentation of the more fragile speci-mens, and partial destruction of the original taphonomic modifications. Thus, the taphono-mic information gained from MFS's is biased towards the compositional features (abun-dance, 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 (some-what more than 44%), isometric (cube-shaped – 19,6%) and irregular (15,6%) ones are also common. The unusual rarity of conical ele-ments is due to the scarcity of isolated teeth.

Hydraulic compatibility between the fine-grained 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 repre-sented 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


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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 cate-gories 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 macro-vertebrates 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 size-related 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

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 well-preserved 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


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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 re-plicated 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 paleo-biocoenosis 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 Fântânele-Vălioara (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


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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 Cam-panian 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 wide-ranging taxon, known from other

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


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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 pre-sence 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 structure-less massive dark gray fine-grained sediments with high frequency of charcoal fragments and pyrite concretions, is also unknown from other sites.

Together, the sedimentological, paleonto-logical 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 com-mon 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 Budu-rone 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 micro-habitat, populating the shore forests, based on the presence of their fragile, but well-preser-ved toothed jaw fragments. The other taxa, such as the terrestrial dinosaurs, were inhabi-tants 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


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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 high-productivity 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 depo-sitional environment, not yet documented by other MFS's: a wet, heavily forested marsh-land 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 semi-aquatic 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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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; 16.2-

143.0; 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.

Z. CSIKI, A. IONESCU & D. GRIGORESCU PLATE I

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Z. CSIKI, A. IONESCU & D. GRIGORESCU PLATE II

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Z. CSIKI, A. IONESCU & D. GRIGORESCU PLATE III

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