-A-D, Novocrania anomala (Müller, 1776), Upper Oligocene; A, B, outer and inner views of ventral valve, Peyrère, MNHN. F.A46481; C, D, inner views of dorsal valves, Abesse, MNHN.F.A46483 and MNHN.F.A46484; E-H, Terebratulina retusa (Linnaeus, 1758), Upper Oligocene, Peyrère; E, dorsal view of young complete specimen, MNHN.F.A46485; F, inner view of dorsal valve to show a brachial skeleton, young individual, SEM, MNHN.F.A46486; G, H, dorsal and lateral views of complete specimen, MNHN.F.A46487. Scale bars: A-D, 2 mm; F, 500 µm; G, H, 5 mm. 

-A-D, Novocrania anomala (Müller, 1776), Upper Oligocene; A, B, outer and inner views of ventral valve, Peyrère, MNHN. F.A46481; C, D, inner views of dorsal valves, Abesse, MNHN.F.A46483 and MNHN.F.A46484; E-H, Terebratulina retusa (Linnaeus, 1758), Upper Oligocene, Peyrère; E, dorsal view of young complete specimen, MNHN.F.A46485; F, inner view of dorsal valve to show a brachial skeleton, young individual, SEM, MNHN.F.A46486; G, H, dorsal and lateral views of complete specimen, MNHN.F.A46487. Scale bars: A-D, 2 mm; F, 500 µm; G, H, 5 mm. 

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Brachiopods from the Upper Oligocene (Chattian), Aquitaine Basin, southwestern France comprise nine species in seven genera: Novocrania, Terebratulina, Megathiris, Argyrotheca, Joania, Megerlia, and Lacazella. One megathyridid is described as a new species, Joania peyrerensis n. sp., characterized by ornamentation of 6-8 distinct, sharp ribs and by...

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... anomala is rare (12 specimens) in the studied material, found only in two localities. e material is poorly preserved, both outer and in- ner surfaces are usually worn but it corresponds well with that hitherto described (Logan 1979(Logan , 1998Logan & Long 2001;Álvarez & Emig 2005). e shell is small with maximum length 15.6 mm, subcircular in outline, wider than long. e posterior adductor muscle scars are large, subcircular, situated near posterior margin in both valves. e anterior adductor muscle scars are oval, elevated. On some specimens mantle canals can be observed (Fig. 2C, D OCCURRENCE. -is species is well known in the Neogene deposits of the Mediterranean province but rare in the Central Paratethys (Dreger 1889; Bitner & Dulai 2004), and it is present in the Aquitaine Middle Miocene (Bitner & Cahuzac 2004). Today it occurs in the north-eastern North Atlantic and the Mediter- ranean Sea (Logan 1979;Brunton & Curry 1979;Curry 1982). Its depth range is from 18 to 2157 m (Logan ...
Context 2
... species has been found in three localities but only in one, Peyrère, it is numerous. e inves- tigated specimens, although often crushed and/ or damaged, are consistent with those hitherto described having, however, smaller size (Logan 1979;Taddei Ruggiero 1985;Gaetani & Saccà 1985;Álvarez & Emig 2005). e shell is ovate to subpentagonal, biconvex, covered with numer- ous, fine ribs that are coarser and strongly beaded in young individuals (Fig. 2E). e cardinalia are typical for the genus with prominent inner socket ridges and deep dental sockets. e loop is preserved only in a young specimen, with crural processes not yet united (Fig. ...
Context 3
... species has been found in three localities but only in one, Peyrère, it is numerous. e inves- tigated specimens, although often crushed and/ or damaged, are consistent with those hitherto described having, however, smaller size (Logan 1979;Taddei Ruggiero 1985;Gaetani & Saccà 1985;Álvarez & Emig 2005). e shell is ovate to subpentagonal, biconvex, covered with numer- ous, fine ribs that are coarser and strongly beaded in young individuals (Fig. 2E). e cardinalia are typical for the genus with prominent inner socket ridges and deep dental sockets. e loop is preserved only in a young specimen, with crural processes not yet united (Fig. ...

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... The crural processes are united, forming a short ring-like loop, peculiar to the genus (Fig. 2H). occurrence: The oldest fossil record of Terebratulina retusa is from the Upper Oligocene of the Aquitaine Basin (Bitner et al., 2013a). This species is well-known and common in the Neogene deposits of the Mediterranean province (Gaetani and Saccà, 1985;Taddei Ruggiero, 1985, 1994Bitner and Moissette, 2003;Koskeridou, 2007;Dulai, 2016Dulai, , 2019Hoffmann et al., 2020) but very rare in the Miocene of the Central Paratethys. ...
Article
Normal 0 21 false false false PL X-NONE X-NONE /* Style Definitions */ table.MsoNormalTable {mso-style-name:Standardowy; mso-tstyle-rowband-size:0; mso-tstyle-colband-size:0; mso-style-noshow:yes; mso-style-priority:99; mso-style-parent:""; mso-padding-alt:0cm 5.4pt 0cm 5.4pt; mso-para-margin-top:0cm; mso-para-margin-right:0cm; mso-para-margin-bottom:8.0pt; mso-para-margin-left:0cm; line-height:107%; mso-pagination:widow-orphan; font-size:11.0pt; font-family:"Calibri",sans-serif; mso-ascii-font-family:Calibri; mso-ascii-theme-font:minor-latin; mso-hansi-font-family:Calibri; mso-hansi-theme-font:minor-latin; mso-bidi-font-family:"Times New Roman"; mso-bidi-theme-font:minor-bidi; mso-font-kerning:1.0pt; mso-ligatures:standardcontextual; mso-fareast-language:EN-US;} Four brachiopod species, i.e., Terebratula cf. styriaca Dreger, Terebratulina retusa (Linnaeus), Megathiris detruncata (Gmelin) and Megerlia truncata (Linnaeus), have been recognised in the Middle Miocene (lower Badenian) deposits at the localities Borač and Borač-Podolí, Carpathian Foredeep, Moravia, Czech Republic. The species M. truncata predominates in the assemblage studied, while M. detruncata is very rare, found only at the locality Borač. Terebratula cf. styriaca and Terebratulina retusa are reported for the first time from the Moravian part of the Carpathian Foredeep. Two types of trace fossils have been observed on the brachiopod shells: drill holes penetrating the shell (ichnogenus Oichnus Bromley) and etching scars, produced by a brachiopod pedicle (ichnogenus Podichnus Bromley and Surlyk).
... Argyrotheca bitnerae was originally described from the Middle Miocene of the Central Paratethys (Dulai and Stachacz, 2011;Hladilová et al., 2014;Dulai, 2015;Kopecká et al., 2018). In Europe, it was also recognized in the Oligocene deposists of France (Bitner et al., 2013a) and Germany (Du-lai and von der Hocht, 2020; Bitner and Müller, 2022). ...
... In the Middle Miocene of the Central Paratethys J. cordata is one of the most common species (Bitner, 1990(Bitner, , 1993Bitner and Pisera, 2000;Bitner and Dulai, 2004;Bitner and Kaim, 2004;Dulai, 2007Dulai, , 2015Zágoršek et al., 2012;Bitner et al., 2013bHladilová et al., 2014;Bitner and Motchurova-Dekova, 2016;Kopecká et al., 2018). Its oldest fossil record is from the Upper Oligocene (Bitner et al., 2013a), and today it lives in the NE Atlantic, Mediterranean Sea and Red Sea, occupying the depth from 3 to 600 m (Logan, 2007;Logan et al., 2008). ...
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The study presents new records of brachiopods from the Lower Miocene deposits of the Qom Formation from two sections, Bagh and Ramsheh in the Isfahan province, Central Iran. Three species, i.e. Argyrotheca bitnerae, Joania cordata, and Platidia anomioides, have been identified in the investigated material. All species are present in the Bagh section whereas in the material from Ramsheh A. bitnerae was not found. Although the species recognized here were already described from the Miocene of the Qom Formation, this is their first report from the localities of Bagh and Ramsheh, providing new data on the distribution of brachiopods in the Cenozoic of Iran.
... The subsequent opening of the Strait of Gibraltar would give rise to the close relationship between the sea pen fauna of Mediterranean and nearby Atlantic regions (Sampaio et al. 2019;Chimienti et al. 2019a). However, the existence of related cnidarian taxa between the Northwestern Mediterranean (Gulf of Lion canyons) and the Southeastern Cantabrian Sea (Gulf of Biscay) may suggest that influence of the Paratethys connexions (throughout the north of the Iberian Peninsula) are still present, and that the possible Tethys relic fauna could have survived the Messiniensis crisis near to main contributions associated with the most important rivers (Rögl 1997;Gili et al. 1998Gili et al. , 2000Bitner et al. 2013;van Baak et al. 2016). ...
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Pennatulaceans (Octocorallia: Pennatulacea) are among the most dominant octocoral species living in soft bottom communities. Some of them are cosmopolitan and have been recorded in all seas and oceans of the world, from shallow waters up to about 6260 m deep. In the present study, we try to find answers to questions about the biogeographical distribution of sea pens including most recently described species. We also evaluate the diversity and faunistic affinities among pennatulaceans of the world's oceans, and determine the degree of faunistic isolation of the Mediterranean Sea from the Atlantic. Finally, we return to the old debate about the hypothetical oldest diversification center of sea pen lineages.
... The poorly preserved, mostly fragmented material and absence of the ventral valve that is taxonomically important (see Robinson, 2017a), prevent any species assignment. The thin shell distinguishes the Ramsheh specimens from the Paleogene and Neogene craniids of Europe (see Dulai, 2004, 2008;Kroh et al., 2008;Bitner et al., 2013;Bitner and Motchurova-Dekova, 2016;Bitner and Müller, 2017). Occurrence: Lower Oligocene Lower Red Formation, Ramsheh section, Iran. ...
... The investigated specimens correspond well, both externally and internally, to those described as Joania peyrerensis by Bitner et al. (2013) from the Late Oligocene of SW France. The specimens from Ramsheh, with the maximum length 2.5 mm, are slightly smaller than those from France. ...
... that live cementing to the substrate by the ventral valve, the remaining species have a functional pedicle opening and live attached to hard substrates. The co-occurrence of thecideide and megathyrid brachiopods is very common, both today and in the fossil record (e.g., Meile and Pajaud, 1971;Bitner et al., 2013;Pedramara et al., 2019). The studied brachiopods indicate a warm, shallow-water environment. ...
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The study presents the first record of Early Oligocene brachiopods from the Lower Red Formation of the Isfahan Province, Central Iran. The assemblage comprises six brachiopod taxa, belonging to five genera, Novocrania Lee and Brunton, Thecidellina Thomson, Argyrotheca Dall, Joania Álvarez, Brunton and Long, and Platidia Costa. Two species are described as new, Thecidellina persica sp. nov. and Argyrotheca ramshehensis sp. nov. The discovery of Thecidellina in Iran is the first confirmed occurrence of this genus in Oligocene deposits, filling the gap between known Eocene and Miocene occurrences. At the species level, the brachiopods investigated here show no affinity to those from the Lower Miocene Qom Formation.
... Occurrence: Lacazella mediterranea has a very long, stratigraphic range with its oldest fossil record from the late Paleocene of Spain (Pajaud and Plaziat, 1972). In the Eocene and Oligocene of Europe, this species is widespread from France to Ukraine (von Koenen, 1894;Bitner and Dieni, 2005;Bitner and Dulai, 2008;Nebelsick et al., 2011;Bitner et al., 2013a;Bitner and Müller, 2017), while in the Neogene its occurrence is restricted to the Mediterranean Province (Logan et al., 2004;Dulai, 2010Dulai, , 2019Hoffmann et al., 2020). Recently, L. medirerranea was identified in the early Miocene of Iran (Pedramara et al., 2019). ...
... Occurrence: Megathiris detruncata is a long ranging species, known since the Eocene (e.g., Bitner and Dulai, 2008;Dulai et al., 2010;Dulai, 2011;Bitner and Müller, 2017). In the Oligocene being known only from few records (Bosquet, 1864;Bitner et al., 2013aBitner et al., , 2020 this study), this species is very common and widespread in the Neogene of Europe (see Gaetani and Saccà, 1985;Bitner, 1990;Bitner and Dulai, 2004;Dulai, 2007;Bitner and Schneider, 2009; Zágoršek et al., 2012;Bitner et al., 2013b;Bitner and Motchurova-Dekova, 2016;Hoffman et al., 2020). Recently M. detruncata was also recognized in the lower Miocene of Iran (Pedramara et al., 2019). ...
... Joania crenata is also distinguished from the late Oligocene species from the Aquitaine Basin, SW France, Joania peyrerensis Bitner, Lozouet and Cahuzac, 2013. In the latter species, the ribs are fewer and extended with large spaces between them, and its median dorsal septum is lower, with four serrrations (Bitner et al., 2013a). Occurrence: This species is known exclusively from the lower Oligocene of Germany (Sandberger, 1862(Sandberger, -1863this study). ...
Article
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Thirteen species, belonging to eleven genera, i.e., Discinisca Dall, Novocrania Lee and Brunton, Lacazella Munier-Chalmas, Pliothyrina van Roy, Terebratulina d’Orbigny, Megathiris d’Orbigny, Argyrotheca Dall, Bronnothyris Popiel-Barczyk and Smirnova, Joania Álvarez, Brunton and Long, Platidia Costa and Megerlia King, have been identified in the lower Oligocene rocky-shore deposits at Mammendorf, central Germany. The occurrence of the species Discinisca fallens (Wood), Argyrotheca bitnerae Dulai and Megerlia truncata (Linnaeus) extends their stratigraphic range back to the early Oligocene. The affinity to the early Oligocene brachiopod fauna of the Mainz Basin is noted.
... However, shortly aft er this publication, some Paleogene records became available. Megerlia truncata was identifi ed from the Chattian (late Oligocene) in France (Bitner et al. 2013), while Megerlia sp. was mentioned also from the Chattian in Hungary (Dulai 2010). Currently the oldest known record is Megerlia sp. from the late Eocene (Priabonian) of Ukraine (Bitner & Müller 2017). ...
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Nannoplankton studies of the Transylvanian Turnu Roșu section, based on the presence of Reticulofenestra umbilicus, R. bisecta, R. lockeri, Helicosphaera compacta and Sphenolithus spiniger refer to Bartonian (middle Eocene) age. The brachiopod assemblage of Turnu Roșu contains four species. The assemblage is dominated by Gryphus kickxii, one of the most common brachiopod species of the European Eocene. The other three taxa are very rare (1-1 specimen). Terebratulina tenuistriata is also widespread in the Tethyan Eocene, while the genus Megerlia everywhere belongs to the very rare elements in the Paleogene. The fourth taxon is an unidentifiable fragmentary terebratulide which is clearly different from the previously mentioned three taxa. The Bartonian Megerlia sp. specimen is the oldest known fossil record of the genus Megerlia and the family Kraussinidae. With 41 figures.
... from the Eocene M. detruncata is reported from Austria, Hungary, and Ukraine(Bitner & Dulai 2008;Dulai et al. 2010;Dulai 2011;Bitner & Müller 2017). Very rare in the Oligocene of Italy (this paper), M. detruncata can be a dominant species in the Oligocene assemblages of France and Hungary(Dulai 2010b;Bitner et al. 2013a). It belongs to one of the most common species in the Miocene of the Central Paratethys (e.g. ...
... The species composition of the studied assemblage differs from the majority of the Palaeogene and Neogene assemblages of Europe where micromorphic megathyridid brachiopods (i.e. Megathiris, Argyrotheca, Joania, and Bronnothyris) dominate (e.g.Davidson 1870;Sacco 1902;Bitner 1990;Dulai 2007;Bitner & Dulai 2008;Bitner et al. 2013a;Bitner & Motchurova-Dekova 2016;Bitner & Müller 2017). Their absence, apart from one specimen of M. detruncata, can be explained by the lack of their preferred environment, hard substrate and cryptic habitats.The Mediterranean species Gryphus vitreus and Terebratulina retusa ...
Article
Four brachiopod species, i.e. Erymnaria bolcensis (Massalongo, 1850), Gryphus cf. minor (Philippi, 1836), Terebratulina tenuistriata (Leymerie, 1846), and Megathiris detruncata (Gmelin, 1791) have been identified in the Oligocene deposits of NW Italy. The species G. cf. minor and T. tenuistriata dominate in the studied material. This is the first record of M. detruncata from the Oligocene of Italy, although it has been noted both from the Eocene and Miocene of Italy. The stratigraphic ranges of E. bolcensis and T. tenuistriata have been extended. The studied assemblage differs considerably from the majority of the Palaeogene and Neogene assemblages of Europe dominated by micromorphic megathyridids. Geological context and sedimentary features point to mass-flow deposits of fan deltas, indicating that the assemblage has been transported within the sediment down a slope.
... Remarks: Terebratulina retusa is a well-known species in the Neogene deposits of the Mediterranean province (Gaetani and Saccà , 1985;Barrier et al., 1992;Encinas and Martinell, 1992;Taddei Ruggiero, 1994;Bitner and Moissette, 2003;García Ramos, 2004;Dulai, 2013Dulai, , 2016Dulai, , 2019. So far its oldest occurrence is reported from the upper Oligocene of the Aquitaine Basin (Bitner et al., 2013a). Today this species lives in the north-eastern Atlantic Ocean and the Mediterranean Sea, with a depth range from 18 to 2157 m (Logan, 2007;Emig, 2018). ...
Article
Marly sediments of the early Messinian Abad Member of the Turre Formation from the northeastern sector of the Carboneras-Nijar Basin (southern Spain) have yielded a rich fossil assemblage, of which 60 taxa are documented herein. Besides nannoflora and microfauna, this assemblage includes the first autochthonous macrofauna described from the Abad Member. Based on the calcareous nannofossil assemblage, in particular the occurrence of the zonal index taxon Amaurolithus primus, the sediments are assigned to the Mediterranean calcareous nannofossil zone CNM17, corresponding to the latest Tortonian to earliest Messinian interval. This matches the age range generally reported for the Abad Member. Palaeoecological evidence from calcareous nannofossils (20 authochtonous taxa), planktic and benthic foraminifera (12 taxa), Porifera (3 taxa), Octocorallia (Keratoisis), Serpulidae (4 taxa), Bivalvia (5 taxa), Gastropoda (2 taxa), Brachiopoda (7 taxa), Cirripedia (Faxelepas) and Vertebrata (5 taxa) indicates an upper bathyal environment with an influx of neritic elements for the Abad Member near Carboneras. Additionally, several faunal components may represent allochthonous/parautochthonous elements from adjacent habitats, which were transported into the deep marine setting by turbiditic mass flows. Although similarities exist, the fossil assemblage from the marls is compositionally significantly different from the biota previously documented from a nearby exposed olistostrome, the ‘red breccia’. Similar fossil assemblages from the Mediterranean have so far mainly been reported from the Pliocene-Pleistocene of southern Italy and Greece. The Carboneras fauna thus adds to our knowledge of the development of these habitats and their biota prior to the Messinian salinity crisis. Beyond the novel palaeoenvironmental data, the range of the dyscoliid brachiopod Ceramisia meneghiniana, previously known only from the Pliocene of Italy, is extended to the Miocene of Spain. The cirripede crustacean Pycnolepas paronai De Alessandri, 1895 is transferred to the hitherto monospecific genus Faxelepas Gale, 2015, whereby the range of the latter (previously Maastrichtian to Danian) is extended to the late Miocene.
... This is compatible with the general ecological niche of M. detruncata based on the modern dataset, but shifted towards its higher end, which is in agreement with the warmer climatic conditions of the late Oligocene (e.g., Zachos et al., 2001). It must be mentioned that Bitner et al. (2013) proposed using δ 18 O seawater of 1 ‰ for their M. detruncata measurements to calculate ambient temperature for the same region. They argue that coastal seawater in the Mediterranean is higher than 1 ‰, hence they assumed similar conditions for the Aquitaine Basin brachiopods. ...
Article
Modern and fossil calcareous shells of the terebratulide brachiopod Megathiris detruncata have been investigated for stable oxygen and carbon isotope and elemental (Mg, Sr, Mn, Fe, Ba) compositions to assess the possibility of using this species to trace past environmental conditions. Modern shells were collected from the Mediterranean Sea and the Atlantic Ocean, while fossil specimens were sampled from late Eocene to Pleistocene deposits in Europe. We mainly analysed bulk shells (i.e., primary and secondary layers together) but, in some cases, their single median and lateral septa, ventral and dorsal valves. The oxygen and carbon isotope compositions of the modern shells indicate that M. detruncata mineralizes the bulk of its calcite shell in equilibrium with seawater. In addition, their Mn, Ba, and Fe concentrations are low with few exceptions from the Atlantic Ocean where high Fe/Ca ratios can probably be related to locally enhanced Fe availability. The Sr/Ca ratio ranges narrowly between 1.4 and 1.7 mmol/mol, while the Mg/Ca ratio has larger variability from about 23 to 45 mmol/mol. The δ18O derived temperatures do not correlate with Mg/Ca, suggesting that Mg is likely affected by greater vital effects. Physiological control on biomineralization might also explain the somewhat larger spread of δ13C between different valves of the same individual and will require further testing by in situ analyses. The fossil M. detruncata shells have a compositional range much larger than modern specimens, with large intra-shell and intra-site variabilities indicative of partial or full alteration. Altered fossils, characterized by low δ18O and δ13C values, low Mg/Ca and Sr/Ca, and high Mn/Ca and Fe/Ca ratios are often associated with clear signs of re-crystallization and/or secondary calcite precipitation within the shells. We used fossil samples without obvious recrystallization features and chemical and isotopic compositions similar to modern specimens to discuss environmental parameters such as temperature and δ18Oseawater at the given time. We recognized variable local controls on seawater isotopic compositions in the Paratethys from the Oligocene until the Middle Miocene, and we propose somewhat warmer habitats for the M. detruncata in the Aquitaine Basin in southwestern France during the Late Oligocene. Isotope variation recorded in the Plio-Pleistocene Mediterranean shells are instead best explained by changing glacial-interglacial conditions.
... The large, hypothyrid foramen is restricted by two narrow deltidial plates. This species is known since the Late Oligocene (Bitner et al. 2013a;Dulai 2010b). It is very common in the Miocene of the Central Paratethys (Bitner 1990 Argyrotheca bitnerae Dulai, 2011. ...
... In the Miocene this species is limited to the northern part of the Central Paratethys (Dulai & Stachacz 2011;Hladilová et al. 2014;Dulai 2015;Kopecká et al. 2018). It was also recognized in the Upper Oligocene deposits of southern France (Bitner et al. 2013a Joania cordata (Risso, 1826) Fig. 6A-G v. 1990 Argyrotheca cordata (Risso, 1826). pl. ...
... The oldest record of this species is from the Late Oligocene of France and Hungary (Bitner et al. 2013a;Dulai 2010b). In the Miocene of the Central Paratethys it belongs to a very common species (Bitner 1990;Bitner & Pisera 2000;Bitner & Dulai 2004;Bitner & Kaim 2004;Dulai 2007;Dulai 2015;Zágoršek et al. 2012;Bitner et al. 2013bHladilová et al. 2014;Bitner & Motchurova-Dekova 2016;Kopecká et al. 2018). ...