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 (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 taphocoenosis made up of locally derived The European continental components that underwent moderate to very assemblages were (and are continuing to be) little transport (except part of the palynoflora). less well known than their Asian, South Consequently, the study of this unique American or especially North American assemblage allows a thorough reconstruction counterparts. This is upheld despite the of a local paleoenvironmental niche inhabited significant advances made in the recent years by the Haţeg continental biota. in the research of the European continental Upper Cretaceous (e.g. Rage, 2002), GEOLOGICAL SETTING especially through continuous fieldwork in The Haţeg Basin is a post-tectonic dep- classical areas such as northern Spain or ression formed in the aftermaths of the Latest southern France (e.g. Le Loeuff, 2001), or in Cretaceous (Second Getic, following Săndu- newly discovered localities such as the lescu, 1984) orogenetic phase of the Alpine Santonian Iharkut locality from Hungary orogenesis, responsible for the thrusting of the (Makádi et al., 2006, Ősi & Rabi, 2006). Our main nappes of the Southern Carpathians. Its in-depth understanding on the Late Creta- formation is linked to local subsidence ceous European continental ecosystems is processes occuring around rapidly uplifting hampered, among other factors, by the almost metamorphic core complexes such as the complete absence of fortunate taphocoenoses Retezat Dome south of the basin, the later yielding both plant and remains, these becoming place of important molasse-type, co-occurrences offering rare insights into the mostly siliciclastic sediment accumulation. composition and structure of the local Thick continental sedimentary rock piles communities (e.g. Calvo et al., 2007) by deposited during the Latest Cretaceous are preserving mostly autochthonous or par- known in the northwestern and central-eastern autochthonous taxa derived from the vicinity of parts of the basin, and were named the the place of their burial. Densuş-Ciula and Sînpetru formations, In the Maastrichtian of the Haţeg Basin respectively (Grigorescu, 1992; Fig. 1). (Southern Carpathians, Romania) only one Although these two units differ somewhat in site yielded both plant and animal remains – lithological composition and overall thickness, the Budurone microvertebrate fossil site (MFS; are usually considered as being time- see Grigorescu et al., 1999). First recognized correlative, a conclusion supported by their for its relatively abundant and diverse largely comparable paleontological content microvertebrates, the special lithology of the (see Grigorescu, 2005, for a recent review). site also allowed the recovery of a similarly Based on age determinations of the underlying diverse plant assemblage, with palynomorphs marine deposits (Grigorescu & Melinte, 2002, and paleocarpological remains. The Melinte & Bojar, 2006, Neagu, 2006), sedimentology of the site, as well as the palynology (Antonescu et al., 1983, van taphonomical features of the local flora and Itterbeeck et al., 2005) and magneto- fauna point to the presence of a stratigraphy (Panaiotu & Panaiotu, 2002),

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]

Z. CSIKI, A. IONESCU & D. GRIGORESCU the deposits are considered of Maastrichtian grey-green micaceous silts and mudstones; age. The presence of the K/T boundary in the sorting is generally poor in these deposits, and sedimentary successions was suggested by coarse-grained terms usually contain high Weishampel et al. (1991), but is not yet proportions of angular or slightly rounded supported by conclusive evidence. clasts derived from a dominantly metamorphic The Budurone MFS is located southwest of source area. Lithoclast petrology, heavy Vălioara village (Fig. 1A), one of the most mineral spectra and sedimentary structures important fossil localities of the Densuş-Ciula within the deposits of the Middle Member Formation, being placed within the fossil- suggest a mainly northern source area, bearing Middle Member of this unit. The MFS represented by the uplifted metamorphic is located in the upstream half of the Budurone basement of the Poiana Ruscă Mts.; minor creek, an ephemeral left-side tributary of the contribution of volcanoclastic material is also Vălioara (or Fântânele) creek that flows from noted in several instances, e.g. in the local north to south through Vălioara village. section of the Tuştea nesting site. Budurone creek was already known to yield Sedimentological observations suggest that vertebrate remains as early as 1915 (Kadic, the deposition of the Middle Member took 1916), Kadic excavating remains from place within a dominantly fluvial setting, several spots (currently impossible to identify) represented by alluvial fans as well as located along the creek. More recently, anastomosed and braided rivers. associated remains of a juvenile individual of The Budurone local section (Fig. 1B), Zalmoxes shqiperorum (Weishampel et al., exposed along the bed and steep walls of the 2003) were recovered from Budurone creek, is represented by gray, gray-green or (Mânzăreanu, 1988, unpubl. B.Sc. thesis), blackish micaceous silts and silty mudstones, apparently somewhat downstream, and at a interbedded with greenish microconglomerates slightly lower stratigraphic level, than the and coarse sandstones. The coarse-grained Budurone MFS discussed in the present beds have an erosional, irregular base cutting contribution. into the underlying finer sediments; they show The fossiliferous potential of the Budurone internal size grading and pass either gradually MFS was first recognized in 1996, being or sharply into the overlying fine ones. The excavated intermittently beginning with 1998, thickness of these beds exceeds several through washing and wet-screening of the centimetres, ranging up to 50 cm. The finer- fine-grained matrix. Altogether, over 800 kg of grained deposits, appearing usually in thicker fossiliferous sediment were processed by beds (up to 1 m), are massive and well standard microvertebrate recovery techniques, consolidated. Their swelling nature suggests and more material is presently being screen- highly smectitic composition, also supported washed. by mass spectrometry analyses (work in The site was mentioned and briefly progress). described by Grigorescu et al. (1999), who The fossiliferous bed is represented by a reported the presence of frog, albanerpetontid, well-sorted, micaceous silty mudstone, over 1 lizard and crocodylian remains. Besides m thick, wedged between two 40-cm thick vertebrates, shells of continental gastropods lithons of coarse sandstones showing graded and carpological remains were also recovered bedding, in the median part of the succession. during the screen-washing process, despite The dark-colored, blackish - slightly bluish silty their fragility and the potentially destructive mudstone is massive, without evident character of the recovery process. Finally, as macroscopic sedimentary structures; it the dark hues of the sediments were contains small, framboidal pyrite concretions suggestive of an environment with high and a large quantity of fusinite (charcoal) potential for palynological material fragments, sometimes up to 0.5-0.7 cm long. preservation, it was sampled and yielded a The sedimentological interpretation of the moderately diverse palynoflora as well. succession suggests the presence of a poorly drained alluvial plain, with areas of high water Lithology and sedimentology of the saturation and ponding, but also influenced by Budurone site fluvial sedimentary processes from nearby The Middle Member of the Densuş-Ciula channels (probably as crevasse splay Formation is made up of repetitive sequences deposits, judging from the sheet-like geometry of alternating coarse- and fine-grained of the coarser beds and lack of identifiable siliciclastic deposits, mainly matrix-supported channel-linked structures within them). conglomerates, lithic sandstones and red and

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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 1.1. Paleocarpology a locally developed small pond or marshy The Budurone MFS is unique within the area, based on the structureless massive cha- Maastrichtian of the Haţeg Basin in yielding racter of the siltstone and the presence of fea- carpological remains. These remains are tures suggesting a reducing geochemistry of rather well preserved, despite their fragility, the depositional micro-environment (charcoal, and the assemblage is relatively diverse. pyrite concretions, plant remains). During the screen-washing of the sediments, several dozens of seeds and fructifications FLORAL AND FAUNAL ASSEMBLAGE OF were recovered, and were sent for study to E.- THE BUDURONE MICROVERTEBRATE M. Friis (Stockholm, Sweden) and her co- FOSSIL SITE workers; the peliminary results of their study shows that the assemblage is interesting. 1. Flora According to these studies (E.-M. Friis, The plant remains are rare in the written comm., 2000; May, 2003, May et al., in Maastrichtian continental beds of the Haţeg prep.), the assemblage includes seeds Basin, especially due to the mostly oxidizing referable to the angiosperm taxon Eurya sp., character of the deposits and to the high as well as fruits of Normapolles type energy of the sediment deposition. Macroplant (Normapolles sp.). The largest part of the remains are especially rare, being confined assemblage is represented by diverse forms mostly to the tuffaceous, fine-grained referred to the carpological metataxon lacustrine deposits of the Lower Member of Carpolithes; about 15 different indeterminate the Densuş-Ciula Formation (see Grigorescu, species of Carpolithes fruit types have been 1992), the main localities being reported identified, including nuts, drupes, seeds and around Densuş village (e.g. Mărgărit & capsules (May, 2003), suggesting a high Mărgărit, 1967). Rare isolated plant fragments, diversity of reproductive strategies already represented by impressions of "Palmophyllum- present in the local Maastrichtian flora. type" angiosperm leaves, are also known from The in-depth study of the Budurone the deposits cropping out along the Sibişel carpological assemblage is yet to be valley at Sînpetru or the Bărbat River valley at undertaken, in order to better understand its Pui. However, plant remains were represented taxic diversity, as well as ecological and up to now only by leaf impressions. evolutionary significance. Palynofloras are better known from these continental deposits (see below); these were 1.2. Palynology recorded from both the Sînpetru and Densuş- The generally reducing character of the Ciula formations and were used to constrain sediments, along with the presence of carpo- the age of the deposits and to reconstruct the logical remains, charred wood and charcoal paleoenvironment the vertebrate fauna lived fragments suggested that the Budurone MFS in. might have good potential for palynomorph 51

Z. CSIKI, A. IONESCU & D. GRIGORESCU

preservation; the sampling of the fossiliferous Cicatricosisporites dorogensis POTONIE & bed yielded indeed a moderately well GELLETICH and Cicatricosisporites sp.), while preserved palynological assemblage. trilete spores dominate (about 80% of the Due to their originally large number and spore assemblage) and include: Appendici- resistance, palynomorphs are easily preserved sporites tricarinatus, Appendicisporites sp., in continental settings. In the Maastrichtian Camarozonosporites insignis NORRIS, Cyathi- continental deposits of the Haţeg Basin, dites australis COUPER, C. sp., Gleicheniidites although more frequently recovered than sp., Cingulatisporites sp., Leiotriletes sp., macroplant remains, their restricted Lycopodiumsporites clavatoides COUPER, L. occurrence is still strongly controlled by the sp., Polypodiaceoisporites sp., Polypodiidites mainly oxidizing character of the sediments. sp., Rouseisporites simplex ROUSE and The first reports on the Latest Cretaceous Stereisporites stereoides PFLUG. Fern spores palynological record from the Haţeg Basin are like Polypodiaceoisporites sp., Polypodiidites that of Stancu et al. (1980) and Antonescu et sp. and Cingulatisporites sp. are dominant, al. (1983). The latter authors identified two being followed by Gleicheniidites sp., relatively rich assemblages from the Densuş- Leiotriletes sp. and bryophytes Ciula Formation (from the neighborhood of (Stereisporites). Vălioara and Densuş, respectively), and a Both gymnosperm and angiosperm pollen somewhat less diverse flora from the Sînpetru was recorded. Gymnosperm pollen is rare Formation at Sînpetru (see also Grigorescu, (about 5% of the assemblage), with Alisporites 1983, 1992). According to this pioneering bilateralis ROUSE and Inaperturopollenites du- study, the assemblage supported a Latest bius PFLUG & THOMSON. Both Normapolles Cretaceous (Late Maastrichtian) age of the (Interpollis, Nudopollis, Occulopollis and deposits. It was dominated by fern spores, but Trudopollis) and post-Normapolles (Subtripo- with a diverse component of angiosperm ropollenites, Triatriopollenites) angiosperm pollen of Normapolles (and subordinately pollen was recovered. The complete list of the Post-Normapolles) type. Another assemblage identified taxa includes: Ephedripites jansonii was recorded, but not published, from the Pui (POCOCK) MULLER, Interporopollenites cf. area by Ştiucă (1983, unpubl. B. Sc. Thesis). nagyae KEDVES & HEGEDUS, I. sp., Interpollis Subsequently, palynological assemblages supplingensis (PFLUG) KRUTZSCH, I. sp., from the Bărbat River (Pui) and Râul Mare Labrapollis sp., Minorpollis sp., Nudopollis cf. River (Toteşti, Nălaţ-Vad) were studied by van terminalis PFLUG & THOMSON, N. thiergartii Itterbeeck et al. (2005). These authors (POTONIE) PFLUG, Occulopollis div. sp., corrected the previous age assignment based Pseudovacuopollis sp., Plicapollis sp., on palynofloras to Maastrichtian, and noted Subtriporopollenites annulatus PFLUG & the important contribution of the gymnosperm THOMSON, Triatrioporopollenites sp. and pollen to the local assemblages, as well as the Trudopollis div. sp. (Plate I, II). dominance of the fern spores and diversity of The Budurone local palynoflora differs from the angiosperm pollen. previously known ones from the Haţeg Basin The samples collected from the Budurone in the importance (both in diversity and abun- MFS were treated by standard palynological dance) of the Normapolles and post-Norma- techniques. The sample was washed and polles angiosperm pollen and from those dried before mechanical disintegration, recorded from the Bărbat and Râul Mare rivers followed by a two-step chemical disintegration (van Itterbeeck et al., 2005) by the reduced with HCl and HF. The separation of the participation of the gymnosperm pollen. These organic material was done through heavy differences probably reflect local environ- liquid separation using ZnCl2. The resulted mental conditions (see below, material was mounted on 10 slides and was Paleoenvironment). studied using an OLYMPUS CX41 transmitted light microscope (all slides are deposited in the 2. Fauna A. Ionescu collections of the Geological 2.1. Invertebrates Institute of Romania). Invertebrates are not uncommonly found in The recovered palynological assemblage the Maastrichtian deposits of the Haţeg Basin, includes only continental taxa, both spores but usually these are represented only by and pollen being well preserved. Spores are indeterminate paper-thin shell fragments; subordinate in abundance, while pollen is screen-washing methodology further contri- dominant both in number and diversity. butes to the destruction of the invertebrate Among the spores (Plate I, II), monolete shells. However, in several instances (inclu- spores are rare (being represented by ding the Budurone MFS) a relatively well pre- Laevigatosporites ovatus WILSON & WEBSTER, served and diverse invertebrate fauna was

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THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT recovered (Antonescu et al., 1983, Pană et al., microvertebrate fossil site from the Haţeg 2002). Basin, where these are exceedingly rare At Budurone, invertebrates are represented (Fântânele – pers. observ.) or absent exclusively by gastropods (Pană et al., 2002). altogether (e.g. Codrea et al., 2002, Smith et The assemblage is dominated by aquatic taxa, al., 2002). Fishes are represented mainly by with diverse lymnaeids: Lymnaea maastrichta lepisosteids (Plate III, figs. 1, 2); both PANĂ 2002 (in Pană, Grigorescu, Csiki et Lepisosteus and Atractosteus can be Costea, 2002), L. dilatata PANĂ, 2002, Physa recognized based on their typical tooth tip patula NICOLAS, 1890, Aplexa galloprovincialis morphology (pointed in Lepisosteus, (MATHERON), 1842, and Eustagnicola cretacea lanceolate in Atractosteus; Gayet, in Sigé et PANĂ, 2002. Besides aquatic taxa, terrestrial al., 1997). A few rhomboidal, thick, ganoid snails are also present, with small sized, but scales known from the site can also be well-preserved cyclophorids [(Rognacia referred to the lepisosteids, a group recorded abreviata (MATHERON), 1832 and Ajkaia cf. from other MFS's in the Haţeg Basin. gregari TAUSCH, 1886)], vertiginids However, the presence of another, non- (Pyramidula grigorescui PANĂ, 2002), lepisosteid fish can also be suggested, based ariophontids (Eoconulus REINHART, 1883) and on the occurrence of an isolated deeply helicids (Arionta precursor PANĂ, 2002 - all amphicoelous fish vertebra, quite unlike the new taxa erected in Pană et al., 2002). opisthocoelous vertebrae of the lepisosteids. The dominance of the aquatic gastropods This vertebra probably belonged to a (currently suggests the presence of a wet environment, indeterminate) teleostean fish (Peng et al., differing from most other sites yielding 2001). significant gastropod faunas in the Anurans Maastrichtian of Haţeg (Pană et al., 2002). Frog remains are outstandingly abundant Besides gastropods, the Budurone MFS at the Budurone MFS. These are being repre- had yielded a large number of small, ovoidal, sented mainly by their characteristic limb ele- longitudinally ribbed and terminally capped ments such as the fused tibiofibulae or radi- structures having a shiny black color. These ulnae. Unfortunately, most of the frog remains structures, once thought to be seeds, are are not diagnostic at a low taxonomic level; currently interpreted as possible insect eggs consequently, the largest part of the Budurone by May (2003). If this inference is confirmed, frog remains indeterminate (Plate III, fig. 5). then the Budurone site will represent the However, Budurone also yielded more second record of Late Cretaceous insects in diagnostic elements as well, including the type the Haţeg paleo-ecosystem (after that specimen of the fossil discoglossid Paralatonia described by Csiki, 2006). transylvanica Venczel & Csiki, 2003 (Plate III, figs. 3-4). It is thus conceivable that at least 2.2. Vertebrates part of the indeterminate frog material belongs Microvertebrates represent an important, also to Paralatonia. However, besides this although only poorly known component of the discoglossid frog the presence of a second Late Cretaceous continental vertebrate faunas anuran taxon can also be suggested based on of Europe. These were only relatively lately a fragmentary maxilla, markedly different from recognized in the Haţeg Basin (e.g. that of Paralatonia. Grigorescu et al., 1985), although the recent Albanerpetonids years witnessed significant additions to our Albanerpetontids, an extinct group of knowledge of the local microvertebrate fauna allocaudatan amphibians (Fox & Naylor, (see Grigorescu et al., 1999, Codrea et al., 1982), are well represented in the Budurone 2002, Smith et al., 2002, Venczel & Csiki, MFS mainly by their peculiar, characteristic 2003, Folie & Codrea, 2005). dentaries with peg-and-socket symphyseal The microvertebrate remains recovered articulation (Plate III, fig. 6); however, isolated, from the Budurone MFS are not abundant fragmentary vertebrae also occur. Fossils (slightly over 100 identifiable elements are referable to albanerpetontids occur frequently known, this number steadily increasing with in the Maastrichtian of the Haţeg Basin (e.g. continuous work on the site), but interestingly, Grigorescu et al., 1999, Codrea et al., 2002, they seem to represent a high diversity Smith et al., 2002, Folie & Codrea, 2005). assemblage with at least 12 different taxa Although formerly (Grigorescu et al., 1999) already recorded, from fishes to . some of these remains were referred to the Fishes Celtedens, this referral was based on The fish remains are relatively rare in the incomplete elements and is now abandoned. Budurone MFS; however, they are better Consequently, all albanerpetontid remains represented here than in any other

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Z. CSIKI, A. IONESCU & D. GRIGORESCU from Budurone can be confidently referred to of crocodylians in the Budurone local Albanerpeton (see also Folie & Codrea, 2005). assemblage. Turtles Theropods Turtle remains are rare at Budurone, Theropods are extremely rare in the despite their wide distribution throughout the Budurone MFS, being represented by only one Maastrichtian continental beds of the Haţeg isolated tooth crown (Plate III, fig. 12). Its Basin. Turtles are represented only by very morphology is similar to that of the tooth genus few small, isolated carapace fragments; their Euronychodon (Antunes & Sigogneau-Russell, surface ornamentation, made up of fine, 1991), previously reported from other MFS's in minute, irregular furrows, is strongly the Haţeg Basin (Csiki & Grigorescu, 1998, reminiscent of that of Kallokibotion bajazidi Codrea et al., 2002), with labially convex and (Nopcsa, 1923), the only chelonian known lingually flat sides. Both the mesial and the from the Haţeg Basin. Consequently, these distal carinae are smooth and placed on the turtle remains are here referred to lingual face of the crown. Kallokibotion. Ornithopods Lizards As in the case of the theropods, ornithopod At Budurone, lizards are uncommon, but remains are rare in the Budurone MFS (only not rare, being represented almost exclusively two isolated shed tooth crowns), although they by fragmentary jaws. The morphological occur frequently in other MFS's, as well as in diversity of the jaws and dentition suggests the macrovertebrate fossil sites or as isolated presence of at least two different lizard taxa. finds. Despite this low frequency, both ornitho- The first, better documented taxon is pod taxa known from the Haţeg Basin (Tel- represented by two incomplete maxillae with matosaurus, Zalmoxes - Plate III, fig. 13; widely spaced, slightly tricuspid, cylindrical Grigorescu, 2005) are represented, based on teeth with pleurodont implantation (Plate III, their very characteristic tooth morphologies figs. 7-8). The overall morphology of the jaw (Weishampel et al., 1993, 2003). fragments suggests scincomorphan affinities. Eggshells A fragmentary maxilla preserving a single Eggshell fragments, not exceeding 3-4 mm almost complete tooth and the largest part of a in maximum dimension, are only uncommonly second one lacking its tip, represents a found in the Budurone MFS. Several second taxon (Plate III, fig. 9). The robust, morphotypes can be identified. One of these barrel-shaped tooth is somewhat reminiscent represents the megaloolithid morphotype, of those of Slavoia described from the Late relatively thick eggshell fragments with the Cretaceous of Asia, and is thus referred here outer surface covered by large tubercles, also to as ?Slavoia. known in the dinosaur eggs reported from Beside these elements, a few isolated several sites spread around the Haţeg Basin vertebrae and toothless jaw fragments are (e.g. Grigorescu et al., 1994, Codrea et al., also referable to indeterminate lizards 2002, Smith et al., 2002). Eggs with this shell Crocodylians morphotype are usually linked to titanosaurian Crocodylian remains appear to be sauropods (e.g. Chiappe et al. 1998), but are exceedingly rare in the Budurone MFS, being considered to belong to the hadrosaur represented only by two isolated teeth; this Telmatosaurus in the Haţeg Basin. Another contrasts markedly with the widespread eggshell morphotype is thinner (not more than presence of the crocodylian teeth in the 1 mm in thickness) and with the outer surface Maastrichtian deposits of Haţeg, either in covered with sparse, crater-like tubercles, macrovertebrate fossil sites, MFS's or as being described from the Upper Cretaceous of isolated finds. southern France as geckonoid eggshells One of these teeth is highly conical, almost (Garcia, 2000). A third morphotype is straight and pointed, with small and smooth represented by thin eggshells, delicately mesial and distal carinae (Plate III, fig. 10). It is ornamented with small tubercles; these might reminiscent of the teeth generally assigned to represent theropod eggshells. the eusuchian Allodaposuchus precedens (Buscalioni et al., 2001; Delfino et al., 2008) TAPHONOMICAL CHARACTERISATION OF and is accordingly referred to this taxon. THE BUDURONE MFS The other tooth is short, widely triangular The taphonomy of the Maastrichtian and labio-lingually compressed (Plate III, fig. continental beds from Haţeg is studied only in 11). It is similar to teeth referred previously by a preliminary manner, the most important Grigorescu et al. (1999) to as Doratodon sp. contributions concerning the characteristic and most probably represents a second taxon macrovertebrate fossil sites, the so-called "fossil pockets" (see Grigorescu, 1983).

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THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT

Recently, a thorough study of the taphonomy sized microvertebrate remains suggests that of several important fossil accumulations was the same agents are responsible for the undertaken by Csiki (2005, unpubl. Ph.D. accumulation of both the sediments and the thesis), and the conclusions of this study will fossils; the same compatibility is obvious in the be published elsewhere; thus, the detailed case of the carpological and invertebrate taphonomical study of the Budurone MFS is remains as well. outside of the scope of this contribution. Taphonomical modifications are repre- The most important taphonomical features sented mainly by fractures; pre-burial and will be, however, discussed shortly, in order to diagenetic fractures as well as fractures gain insights into the nature of the original related to the recovery process are all present depositional environment and the potential and sometimes difficult to separate. Bone taphonomical biases that operated. splinters are frequent. Weathering and The taphonomical study of the MFS's abrasion of the microvertebrate remains is (including Budurone) requires a specific variable; while the better preserved, approach, due to the recovery method of the taxonomically determinate specimens are only fossils, through bulk sediment sampling and slightly weathered and abraded, if at all, the screen-washing. The recovery procedure indeterminate bone chips and splinters are implies first of all the almost complete loss of usually heavily weathered so as to expose the information concerning the spatial distribution, internal bone structure and are also highly orientation or possible association/articulation abraded. The weathering and abrasion stages of the different specimens. Moreover, the reco- shown by the bone fragments suggest these very procedure is destructive, leading to experienced repeated episodes of transport further fragmentation of the more fragile speci- and reworking. This was not, however, the mens, and partial destruction of the original case of the better-preserved remains. Besides taphonomic modifications. Thus, the taphono- being significantly less abraded and mic information gained from MFS's is biased weathered (showing the unaltered smooth towards the compositional features (abun- shiny outer cortex of the bones), these also dance, diversity) and against the detailed include extremely fragile elements such as understanding of the taphonomic processes tooth-bearing jaws fragments of amphibians or operating. lizards. The above observations suggest that Taking into account this bias, a good this component of the thanatocoenosis amount of taphonomic information is still experienced either only very short transport recoverable, allowing the reconstruction of the and short-term subaerial exposure or none at major aspects of the local assemblage's all (excepting possibly the rare isolated teeth taphonomic history. that, being among the most resistant elements Taphonomic characteristics and of the skeleton, can better withstand transport modifications and subaerial exposure). Several hundred microvertebrate remains Composition, abundance and diversity were recovered; however, the largest part of Although based only a limited sample these are indeterminate bone fragments, while (slightly more than 100 specimens), the only slightly over 100 specimens are vertebrate assemblage preserved in the determinable at least at major clade (e.g. Budurone MFS is relatively diverse, as it anuran, lizard) level. The microvertebrate includes at least 12 different vertebrate taxa. remains and associated flora and fauna seem The most diverse group is that of the fishes, to be concentrated into the upper half of the with 3 different taxa, which is outstanding fossiliferous bed. compared to other Haţeg local assemblages in The bone assemblage shows a low which fish remains are extremely rare or dimensional variability (many specimens have absent altogether. Ornithopods, crocodylians, maximum dimensions of less than 1 mm, and anurans and lizards are slightly less diverse only a minor percent reaches a maximum (each being represented by 2 taxa – maybe 3, diameter of over 5 mm); thus, size sorting in the case of the frogs), while theropods, seems to be good to excellent. Morphological turtles and albanerpetontids are the least diversity is, by the contrary, rather high; diverse clades, with only one taxon present. although long tubular bones dominate (some- The taxic abundance is estimated based on what more than 44%), isometric (cube-shaped the minimum number of identified specimens, – 19,6%) and irregular (15,6%) ones are also as proposed previously for microvertebrate common. The unusual rarity of conical ele- sites consisting mainly of disarticulated and ments is due to the scarcity of isolated teeth. isolated remains (e.g. Foster, 2001, 2003); this Hydraulic compatibility between the fine- approach assumes that each element grained sediments and the dominantly small- represents another individual, as skeletal

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Z. CSIKI, A. IONESCU & D. GRIGORESCU association cannot be proved. Based on this microvertebrate remains, but of its special count, anurans are the most abundant group taphonomic signature. (about 75% of the identifiable remains), The most remarkable taphonomic followed by the lizards (10%), albanerpetontids characteristics of the Budurone MFS are (5%) and fishes (4%). On the other end of the represented by the: a) quasi-absence of abundance spectrum, the crocodylians, subaerial weathering and abrasion in the theropods, ornithopods and turtles are rare taxonomically identifiable elements; b) good (under 2% NIS). preservation state of fragile skeletal elements However, confronting the abundance and such as toothed jaw fragments, diversity data shows some interesting albanerpetontid vertebrae, but also the patterns. While the material referable to the angiosperm seeds and fruits or the relatively two (or possibly three) anuran taxa totals well preserved thin-shelled gastropods etc.; c) about three-quarters of the NIS, the fishes, low frequency of the otherwise resistant with a comparable taxic diversity figure, skeletal elements, commonly recovered from yielded only 4 specimens (less than 4%). other MFS's from Haţeg, such as isolated Similarly, the taxonomically least diverse teeth or turtle plates; d) high albanerpetontids are represented by several frequency of indeterminable bone fragments specimens (5%), but the relatively more and scraps showing complex fracturing, as diverse crocodylians or ornithopods are well as advanced weathering and abrasion. identified based on very few remains (less Together, these features suggest that the than 2%). Budurone assemblage represents a composite All major skeletal element categories (skull attritional taphocoenosis, including two fragments, isolated teeth, vertebrae, girdle, different parts – one with a high degree of limb and dermal elements) are represented, taphonomical elaboration (represented by the although the elements of the appendicular bone fragments) and one with low degree of skeleton dominate by far (over 44%); this is taphonomical alteration (measured either by mainly due to the over-representation of the weathering, abrasion or fracturing). This anuran tibiofibulae, humeri and phalanges. second component most probably represents Jaw fragmens and vertebrae are also relatively a locally derived, autochthonous or common (16,7%), while other cranial elements parautochthonous assemblage, while the (1%) and girdles (2%) are rare. Especially taphonomically altered component is noteworthy is the extreme rarity of the isolated allochthonous, and probably reworked in teeth (of and crocodylians) and several phases. Consequently, the well- dermal elements (reduced to turtle shell preserved component of the taphocoenosis fragments); both of these categories occur (and the one that includes the largest part of commonly in other MFS's. the taxonomically identifiable elements) Distribution of the different skeletal cate- reflects a local biocoenosis, with a limited gories among the major taxonomic groups amount of spatial and time averaging. represented shows a pattern that occurs The assemblage is politaxic with a large frequently in other MFS's in the Haţeg area: number of individuals represented, and typical microvertebrates such as anurans, and taxonomically relatively diverse. However, it to a lesser extant albanerpetontids and lizards shows some interesting pattern such as: (as well as the otherwise almost unknown a) low intra-clade diversity (between 1 and fishes) are represented by a wide range of maximum 3 taxa in each group, including skeletal elements from fragile cranial to more otherwise diverse clades such as lizards, robust vertebral elements, while macro- theropods or crocodilians); vertebrates are known almost exclusively by a b) several compositional peculiarities, single skeletal category: isolated teeth compared to other Haţeg MFS's, such as the (crocodylians, dinosaurs) or dermal plates extreme abundance of the frog remains, (turtles). In this last case, the categories relative high diversity of the fishes (otherwise involved include the most resistant elements of almost unknown from other MFS's), as well as the skeleton, suggesting that macrovertebrate the rarity of turtle, crocodylian and dinosaur remains underwent specific taphonomic remains (usually more frequently found in the processes selecting (besides a normal size- different MFS's) related bias) for those remains that best c) a distinction between two different withstand weathering, breakage and abrasion. categories of the identifiable remains, based Discussions on skeletal representation. The first category The Budurone MFS is among the most (A), illustrated first of all by frogs, but to a important ones in the Haţeg Basin, mainly not lesser extent also by albanerpetontids, fishes because of the quantity of the recovered and lizards, is represented by taxa with

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THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT numerous and diverse skeletal elements Grigorescu, 2005), although discovery of new preserved, belonging to different parts of the taxa (especially invertebrates, plants, but also body and including extremely fragile skeletal vertebrates) is to be expected. Different sites parts in good preservation state. The second yielded local assemblages with slightly category (B) includes the turtles and different compositions, but usually the nature archosaurs, represented by few elements from of the main groups represented and their only one skeletal category, usually one with relative participation are largely predictable. high preservation potential. The Budurone MFS, however, yielded a rather d) the differential preservation is also re- peculiar floral and faunal assemblage, with plicated by differences in size (category A taxonomic composition and especially relative includes small-sized taxa, while category B abundance of the taxa that set it apart from larger-sized ones) and ecological preferences other sites of the Haţeg area. (category A are mainly strictly aquatic taxa, Fishes are extremely rare in the Haţeg while category B mainly strictly terrestrial, or fauna, besides Budurone being reported from semiaquatic, but not water-bound ones). only two other MFS's: from the Pui locality This twofold division of the identifiable excavated by a Romanian-French team remains in the Budurone assemblage (Grigorescu et al., 1985) and from Fântânele- suggests that it is formed by two distinct Vălioara (Grigorescu et al., 1999). At Pui, biocoenotic components. One is locally acipenseriform and characid fish remains were derived, autochthonous, with no or extremely found, while Fântânele yielded lepisosteid and reduced transport from the microhabitat it possible characid remains. The rarity of fish actually inhabited, the skeletal remains being remains in Haţeg contrasts markedly with their buried in large numbers shortly after the death common occurrence in other European Late of the organisms, within their living Cretaceous faunas where lepisosteids are environment. This component also includes, common, and usually extremely abundant, besides most of the category A vertebrate while other taxa are also present (e.g. Cavin, remains (excepting possibly the lizards), also 1999, Makádi et al., 2006). In this respect, the the carpological remains and gastropods as Budurone MFS is outstanding in showing the well. The other component is a highest fish diversity with three different taxa, parautochthonous one, derived from nearby although their remains are rare. microhabitats, but without much reworking or Anuran remains occur frequently in the long-distance transport; occasional sediment different MFS's around the Haţeg Basin (e.g. influx is probably responsible for the Grigorescu et al., 1999, Codrea et al., 2002, introduction of some of these components, as Smith et al, 2002, Folie & Codrea, 2005). suggested by the hydraulic compatibility Although usually not identifiable to lower between them and the surrounding taxonomic level, their very characteristic sedimentary matrix. However, selection of the remains are easily recognizable and thus allow remains occurred during this transport by the identification of the clade. Most anuran preferential concentration of the more resistant remains are referred to different taxa within the elements such as isolated teeth or shell Discoglossidae (Grigorescu et al., 1999, fragments (category B elements). One Venczel & Csiki, 2003, Folie & Codrea, 2005), exception to this is represented by the lizard but other taxa were also reported (Venczel & remains, included in category A, but derived Csiki, 2003). At Budurone, several anuran from strictly terrestrial taxa, suggesting the remains are referable to the Discoglossidae, lizards inhabited closely located microhabitats and some of them to the discoglossid and underwent extremely short transport, Paralatonia transylvanica (Venczel & Csiki, while other terrestrial taxa lived farther from 2003), the type locality of which is represented the site of the burial. The semi-aquatic turtles by Budurone. Paralatonia is also known from and crocodilians, rare in the Budurone MFS, other MFS's from the Haţeg Basin. The taxon despite their common occurrence in other is not yet reported from outside the Haţeg MFS’s, represent another partial exception. area. Besides Paralatonia, another anuran This is probably explained by the fact that the taxon is also present; it is rare at Budurone, specific depositional environment at Budurone, and not yet discovered in other MFS's. although wet, was not particularly suitable for Fossils referable to the extinct group of these taxa. allocaudatan albanerpetontids occur frequently in the Maastrichtian of the Haţeg Basin (e.g. DISCUSSIONS Grigorescu et al., 1999, Codrea et al., 2002, Faunal composition of the Budurone MFS Smith et al., 2002, Folie & Codrea, 2005). The composition of the Haţeg paleo- Although formerly (Grigorescu et al., 1999) biocoenosis is now relatively well-known (see some of these remains were referred to the

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Z. CSIKI, A. IONESCU & D. GRIGORESCU genus Celtedens (now known to be restricted Transylvanian sites outside Haţeg (Codrea & to the Early Cretaceous, McGowan, 2002), this Dica, 2005; Delfino et al., 2008), but also from referral was based on incomplete elements southern France and Spain (Buscalioni et al., and are now abandoned. Consequently, all 2001). Doratodon is also known outside albanerpetontid remains can now be Romania, from the Santonian of Hungary (Ősi confidently referred to Albanerpeton (see also & Rabi, 2006), as well as the Campanian of Folie & Codrea, 2005), a genus with a wide Austria (Seeley, 1881) and Spain (Company et stratigraphic distribution, ranging from the al., 2005). Aptian to the Pliocene (Gardner, 2002, The rarity of dinosaur remains in the Gardner et al., 2003, Venczel & Gardner, Budurone assemblage, reduced to but one 2005). Albanerpetontids, mostly indeterminate isolated tooth for each taxon represented, is taxonomically, are reported from the again remarkable, compared to other MFS's Santonian of Hungary (Makádi et al., 2006) as from the Haţeg Basin. Euronychodon is a well as from several western European Late small theropod, known only by its distinctive Senonian localities (e.g. Buffetaut et al., 1999, teeth, being recovered from several sites in Duffaud & Rage, 1999, Garcia et al., 2000). Haţeg (Csiki & Grigorescu, 1998, Codrea et The turtle Kallokibotion is frequently al., 2002). It is also known from several occurring in the fossiliferous sites in the Haţeg western European localities, both from the Basin, and it also present in the Budurone Lower (Rauhut, 2002) and the Upper MFS. However, its remains are unusually rare Cretaceous (Antunes & Sigogneau-Russell, here, while they are common in most MFS's, 1991) of Iberia. However, the absence of as well as macrovertebrate sites. Kallokibotion dromaeosaurid teeth, common in other sites, is not yet reliably reported from the Late is surprising. Cretaceous of Europe outside Romania On the other hand both Zalmoxes and (Gaffney & Meylan, 1992, Codrea & Dica, Telmatosaurus, the ornithischians usually 2005), except maybe from the Lower Cam- represented in other fossiliferous sites from panian of Austria (Sachs & Hornung, 2006). the Haţeg Basin, including MFS's as well, are Lizards are a recently discovered present at Budurone. They are among the component of the Haţeg local fauna, being first most common dinosaurian taxa from the reported by Grigorescu et al. (1985), but Haţeg Basin and surrounding areas (pers. described on some detail only by Grigorescu obs.; Codrea & Dica, 2005). Zalmoxes is et al. (1999). Since then, several important probably endemic to the Transylvanian area, contributions were published (Codrea et al., but is possibly also present in the Lower 2002, Smith et al., 2002, Folie & Codrea, Campanian of Austria (Sachs & Hornung, 2005) revealing a diverse lizard component of 2006); Telmatosaurus, however, seems to be the assemblage. Lizard remains are commonly restricted to the Transylvanian area and occurring in different microvertebrate fossil unknown from more western parts of Europe sites (Grigorescu et al., 1999, Codrea et al., (Laurent et al., 1997). 2002, Smith et al., 2002, Folie & Codrea, In overall, the composition of the Budurone 2005), but few taxa were discovered in more local assemblage is comparable to that of than one site. The local diversity of the lizards other Haţeg MFS's, but differs from all of these rarely exceeds two to four taxa, this being the in the presence of a diverse fish component, in case also at Budurone. One of the taxa the rarity of the terrestrial archosaurs and represented is unknown from other MFS's, overwhelming abundance of the frogs. while the other one (?Slavoia) is probably Moreover, it is as yet unmatched in that present at Pui as well. None of these taxa invertebrates and especially plant remains were previously reported from other Late (palynomorphs, fruits and seeds) are Cretaceous European localities, although our associated to the microvertebrates. This knowledge both about the lizards from Haţeg unique composition is probably due to the and Europe is still admittedly poor. unique microhabitat represented by the Both Allodaposuchus and Doratodon occur depositional environment of the fossiliferous frequently in different MFS's in Haţeg. bed. Allodaposuchus is rather common also in Biostratigraphy macrovertebrate sites or as isolated finds, From the Budurone local assemblage, the while Doratodon, probably a smaller-sized palynomorphs are especially useful as age taxon based on its known remains, is more indicators. Most of the recorded palynomorph rarely found in these circumstances. Their taxa have a Late Cretaceous-Early Tertiay remains are, however, surprisingly rare in the range, but typical Paleocene taxa are missing. Budurone MFS. Allodaposuchus is a wide- The dominance and diversity of the ranging taxon, known from other Normapolles group, especially the presence of

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THE BUDURONE MICROVERTEBRATE SITE FROM THE MAASTRICHTIAN OF THE HAŢEG BASIN - FLORA, FAUNA, TAPHONOMY AND PALEOENVIRONMENT such taxa as Interpollis, Nudopollis, and tree ferns, a vegetation different both from Occulopollis and Trudopollis, suggest a the mangrove vegetation reconstructed for the Maastrichtian age for these deposits (Goczan nearby Rusca Montană Basin (Petrescu & et al., 1967, Batten, 1981, Portiagina, 1981, Duşa, 1980) or the more open, wooded Herngreen et al., 1986), similar to those savannah vegetation of the Râul Mare-Râul studied by van Itterbeeck et al. (2005). Bărbat area (van Itterbeeck et al., 2005). Paleoenvironment Paleoecology Several lines of evidence point to the pre- Taphonomical observations on the Budu- sence of a particular depositional environment rone microvertebrate assemblage suggest the at Budurone, one that differs from those mixture of two different paleocommunities – represented by other MFS's. These features one autochthonous to the depositional include: the dominance of aquatic gastropods, environment and one parautochthonous, the preservation and high diversity of derived from nearby areas. The angiosperm seeds and fruits (duplicated by the autochthonous paleobiocenosis is represented high diversity and dominance of angiosperm mainly by aquatic and semi-aquatic taxa, such pollen), the dominance of the remains of as fishes, frogs and albanerpetontids, aquatic-semiaquatic taxa (especially frogs), inhabiting the pond or swamp and its thickly the unusual diversity of fishes, the rarity of vegetated shore forests. Based on the iliac typical terrestrial taxa, represented exclusively morphology, Paralatonia was considered to be by remains with high preservation potential a frog with good swimming and/or jumping and the predominantly local derivation abilities, inhabiting riverine or marsh (autochthony) of the microvertebrate environments (Venczel & Csiki, 2003). thanatocoenosis, as suggested by the Albanerpetontids, more terrestrial in habitat, taphonomic features. The sedimentological were showed to live in densely forested areas character complex, especially the structure- as well (Venczel & Gardner, 2005). Lizards, less massive dark gray fine-grained sediments although terrestrial in habitat preference, might with high frequency of charcoal fragments and have also formed elements of this micro- pyrite concretions, is also unknown from other habitat, populating the shore forests, based on sites. the presence of their fragile, but well-preser- Together, the sedimentological, paleonto- ved toothed jaw fragments. The other taxa, logical and taphonomical evidence points the such as the terrestrial dinosaurs, were inhabi- presence of a wet, water-logged or ponded tants of nearby drier, more terrestrial habitats, environment, probably a small swamp or but probably ventured into the forested shores shallow, marshy pond microhabitat within a of the pond either for browsing or hunting. poorly drained floodplain. The rarity of crocodylian and turtle remains The lack of any dinoflagellate cysts – both semi-aquatic – is somehow puzzling, suggests a typical continental setting for the and probably means that these taxa were not Budurone depositional site, weakening the perfectly adapted to the local conditions and proposed arguments for the presence of favored more open, lacustrine or fluvial marine or brackish conditions in the Vălioara settings. The absence of the multituberculates, area (e.g. Vremir, 1998). occurring in many other MFS's, is also The palynological assemblage points to the unexpected, especially since the high diversity presence of a warm, tropical-subtropical of angiosperms would have represented a rich climate, suggested by the rarity of bisaccate (3 food source to these taxa. The small number Alisporites specimens) and taxodiacean (one of identifiable remains might account for their Inaperturopollenites species) pollen, the com- apparent absence, or, alternatively, they as mon presence of ferns such as Gleicheniidites, well would have preferred drier habitats. Lygodiumsporites, Cicatricosisporites and The image of a complex local food web is Cyathidites. The presence of a somewhat drier emerging from the study of the Budurone subtropical climate is also supported by the MFS. The base of the web is represented by a abundance of the Normapolles pollen group rich angiosperm assemblage, completed by (Pacltova, 1966, Zaklinskaya, 1977). water-loving ferns and bryophytes. The The high diversity of Normapolles pollen, vegetation represented the food source for but especially that of the angiosperm seeds invertebrates and probably for herbivorous and fruits (whose good preservation, despite dinosaurs entering this pond-shore forest from their fragility, suggests short transport and their preferred drier habitats nearby. The thus autochthony) allows the reconstruction of preserved autochthonous invertebrates, the local vegetation as being closed, forested, gastropods and possibly insects, were forming made of a thick angiosperm ground cover and a link between the primary producers and low canopy, with participation of herbaceous secondary invertivorous consumers such as

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Z. CSIKI, A. IONESCU & D. GRIGORESCU lizards, frogs, albanerpetontids and possibly rendus Académie des Sciences de Paris, II/313, some fishes. The top of the local food web 113-119. was represented by predatory fishes such as Batten, D. J., 1981. Stratigraphic, paleogeographic lepisosteids, along with rare and and evolutionary significance of Late Cretaceous and Early Tertiary Normapolles parautochthonous crocodylians and Pollen. Review of Palaeobotany and theropods, visiting intermittently this high- Palynology, 35, 125-137. productivity and biomass-rich area. Buffetaut, E., Le Loeuff, J., Tong, H., Duffaud, S., Cavin, L., Garcia, G., Ward, D. & l’Association CONCLUSIONS culturelle, archeologique et paleontologique de The discovery of the Budurone MFS adds Cruzy, 1999. Un nouveau gisement de important information to our knowledge about vertébrés du Crétacé supérieur a Cruzy the Maastrichtian continental ecosystem of the (Hérault, Sud de la France). Comptes Rendus Haţeg Basin. It represents a particular depo- de lAcadémie des Sciences Paris, 328, 203- 208. sitional environment, not yet documented by Buscalioni, A. D., Ortega, F., Weishampel, D. B. & other MFS's: a wet, heavily forested marsh- Jianu, C. M., 2001. A revision of the land or swampy pond preserving a good crocodyliform Allodaposuchus precedens from sample of the local and more distal the Upper Cretaceous of the Haţeg Basin, paleobiocoenoses. The local paleobiocoenosis Romania. Its relevance in the phylogeny of included a rich angiosperm assemblage, Eusuchia. Journal of Vertebrate Paleontology, documented by pollen, but also by seeds and 21, 1, 74-86. fruits, extremely rarely preserved in the Late Calvo, J. O., Porfiri, J. D., Gonzáles-Riga, B. J. & Cretaceous fossil record and otherwise as yet Kellner, A. W. A.. 2007. A new Cretaceous terrestrial ecosystem from Gondwana with the unknown in the Haţeg area. This type of floral description of a new sauropod dinosaur. Anais assemblage differs from those previously da Academia Brasileira de Ciências, 79, 3, 529- reconstructed, suggesting the presence of a 541. complex, mosaic-like vegetation pattern. The Cavin, L., 1999. Osteichthyes from the Upper Budurone local assemblage also supports Cretaceous of Lano (Iberian Peninsula). Est. previous paleoclimatic and biostratigraphic Mus. Cienc. Nat. de Alava, 14, 105-110. interpretations. The microvertebrates, together Chiappe, L. M., Coria, R. A., Dingus, L., Jackson, with the invertebrates and plants, allow the F., Chinsamy, A. & Fox, M., 1998. Sauropod reconstruction of a complex local food web, dinosaur embryos from the Late Cretaceous of Patagonia. Nature, 396, 258–261. dominated by small-sized aquatic and semi- Codrea, V., Smith, T., Dica, P., Folie, A., Garcia, G., aquatic invertivores. Godefroit, P. & Van Itterbeeck, J., 2002. Dinosaur egg nests, mammals and other ACKNOWLEDGMENTS vertebrates from a new Maastrichtian site of the The authors wish to thank all the Haţeg Basin (Romania). Comptes Rendus participants of the Haţeg field campaigns who Palevol, 1, 3, 173-180. contributed to the excavation of the Budurone Codrea, V. & Dica, E. P., 2005. Upper Cretaceous - MFS and sorting of the specimens. Dr. M. lowermost Miocene lithostratigraphic units Venczel helped in the identification of the exposed in Alba Iulia – Sebeş – Vinţu de Jos area (SW Transylvanian basin). Studia anurans, albanerpetontids and lizards; his Universitatis Babeş-Bolyai, Geologia, 50, 1-2, drawings are reproduced in Plate III. The SEM 19-26. photographs were taken in Bucharest (by Dr. Company, J., Pereda Suberbiola, X., Ruiz- C. Costea) and Bruxelles (Julien Cillis). The Omenaca, J. I. & Buscalioni, A. D., 2005. A new suggestions of the anonymous reviewers are species of Doratodon (: also thanked. The fieldwork was supported by ) from the Late Cretaceous of the CNCSIS grants G239/1998-2001 and Spain. Journal of Vertebrate Paleontology, 25, G1163/2004-2006, as well as EC FP5 Project 2, 343–353. ABC 95/2003. Csiki, Z., 2006. Insect borings in dinosaur bones from the Maastrichtian of the Haţeg Basin,

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Albanerpetontid amphibians Ecosystems, Acta Palaeontologica Polonica, 28, from the Lower Cretaceous of Spain and Italy: a 1-2, 103-121. description and reconsideration of their Grigorescu, D., 1992. Nonmarine Cretaceous systematics. Zoological Journal of the Linnean Formations of Romania. Chen, P.-J. & Matter, Society, 135, 1–32. N. J. (eds.), Aspects of Nonmarine Cretaceous Melinte, M. & Bojar, A.-V., 2006. Upper Cretaceous Geology, Beijing, China Ocean Press, 142-164. marine red beds in the Haţeg area (SW Grigorescu, D., 2005. Rediscovery of a “forgotten Romania). In Csiki, Z. (ed.), Mesozoic and land”. The last three decades of research on the Cenozoic Vertebrates and Paleoenvironments.

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Contribuţii la studiul Palaeobotanist, 15. 52-54. petrografic şi paleontologic al Depresiunii Haţeg Panaiotu, C. & Panaiotu, C., 2002. Paleomagnetic pe baza unor foraje. D. S. Inst. Geol. Geofiz., studies. In Grigorescu, D. C. & Csiki, Z. (Eds.), 67, 4, 115-136. 7th European Workshop on Vertebrate van Itterbeeck, J., Markevich, V. S. & Codrea, V., Palaeontology, Abstracts volume and 2005. Palynostratigraphy of the Maastrichtian excursions field guide. Ars Docendi, Bucharest, dinosaur and mammal sites of the Râul Mare 59. and Bărbat valleys (Haţeg Basin, Romania). Pană, I., Grigorescu, D., Csiki, Z. & Costea, C. Geologica Carpathica, 56, 2, 137-147. (2002). Paleo-ecological significance of the Venczel, M. & Csiki, Z., 2003. New discoglossid continental gastropod assemblages from the frogs from the Latest Cretaceous of Haţeg Basin Maastrichtian dinosaur beds of the Hateg Basin. (Romania). Acta Palaeontologica Polonica, 48, Acta Palaeontologica Romaniae, 3, 337-343. 4, 599-606. Peng, J. H., Russell, A. P. & Brinkman, D. B., 2001. Venczel, M. & Gardner, J. D., 2005. The Vertebrate microsite assemblages (exclusive of geologically youngest albanerpetontid Mammalia) from the Foremost and Oldman amphibian, from the Lower Pliocene of Hungary. formations of the Judith River Group Palaeontology, 48, 6, 1273-1300. (Campanian) of southeasern Alberta: An Vremir, M., 1998. The presence of an advanced sea illustrated guide. The Provincial Museum of turtle (Testudines: Chelonioidae) in the Late Alberta, Natural History Occasional Paper, 25, Cretaceous Sînpetru Formation of Haţeg Basin. 1-54. Abstracts volume, Second Symposium on Petrescu, I. & Duşa, A. 1980. Flora din Cretacicul Mesozoic Vertebrate Faunas of Central Europe, superior de la Rusca Montană – o raritate în Deva. patrimoniul paleobotanic naţional. Ocrotirea Weishampel, D. B., Grigorescu, D. & Norman, D. Naturii şi a Mediului Înconjurator, 24, 147-155. B., 1991. The Dinosaurs of Transylvania. Portniagina, L. A., 1981. 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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; 16.2- 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.

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

64

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

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

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