— Aphelesia bipartita (Brocchi, 1814); A-H, ventral, dorsal, lateral and anterior views of complete specimens, Draria, Algeria (LP-MNHN B.38645); I-L, ventral, dorsal, lateral and anterior views of complete specimen, Mustapha supérieur (Algiers), Algeria (LP- MNHN B.38648); M, interior view of dorsal valve, Bab El Oued, Algeria (LP-MNHN B.38658). Scale bars: 1 cm.  

— Aphelesia bipartita (Brocchi, 1814); A-H, ventral, dorsal, lateral and anterior views of complete specimens, Draria, Algeria (LP-MNHN B.38645); I-L, ventral, dorsal, lateral and anterior views of complete specimen, Mustapha supérieur (Algiers), Algeria (LP- MNHN B.38648); M, interior view of dorsal valve, Bab El Oued, Algeria (LP-MNHN B.38658). Scale bars: 1 cm.  

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2003. — Pliocene brachiopods from north-western Africa. Geodiversitas 25 (3) : 463-479. ABSTRACT Seven species of Pliocene brachiopods belonging to the genera Aphelesia Cooper, 1959, Terebratula Müller, 1776, Gryphus Megerle von Mühlfeld, 1811, Terebratulina d'Orbigny, 1847, Megathiris d'Orbigny, 1847 and Megerlia King, 1850 have been recognised in...

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... 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. So far, it has been recorded from Austria (Dreger, 1889), Hungary (Bitner and Dulai, 2004) and the Czech Republic (this study). ...
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... This taxon is profusely recorded in the Upper Miocene-Lower Pliocene of the Mediterranean basins. In the westernmost peri-Iberian basins (Mediterranean-Atlantic seaway), Aphelesia is recorded in the Late Tortonian, and re-emerged again in the earliest Pliocene (Bitner and Martinell, 2001;Bitner and Moissette, 2003;García-Ramos, 2004;Toscano-Grande et al., 2010;Giannetti et al., 2018) when the background conditions after the Messinian Mediterranean Salinity Crisis were re-established (Giannetti et al., 2018). ...
Article
Genetic control undoubtedly plays the most decisive role in the development of characters in rhynchonellides, especially in those structural features such as shell microstructure and brachidial architecture. However, considering the relationship between shell microstructure and type of crura in post-Palaeozoic rhynchonellides supported by recent growing research, the present study links this striking matching to the distribution of rhynchonellide morphogroups around the main post-Palaeozoic mass extinction and diversification events, innovatively introducing the ribbing pattern as a feature widely argued as indicator of deeper/shallower habitats and particular ecological conditions. The analyses performed on the Western Tethys rhynchonellide database and, directly on 28 species from the peri-Iberian margins around selected post-Palaeozoic extinction/diversification episodes, reveal a recurrent distribution pattern according to the habitat considered (epicontinental vs. epioceanic). The smooth, eurinoid, septifal/arcual body plan is the distinctive inhabitant of deeper epioceanic environments, also being the last to withdraw from the epicontinental platforms when the environmental conditions were adverse, and the pioneers or opportunistic taxa conducting repopulation of these epicontinental habitats when the conditions were re-established after crises. The ribbed, eurinoid, falciform morphogroup seems to be the more resilient stock on epicontinental platforms in relation to ecological crises. Conversely, the ribbed, leptinoid, raducal group is typical inhabitant of epicontinental seas when normal conditions prevail and shows an uneven distribution unlinked to critical events, suggesting a more specialist pattern with a diversification related with a secondary differentiation. It is suggested that this distribution dynamic of morphogroups was controlled by the availability of requirements to generate each type of crura and microstructure in epicontinental/epioceanic habitats, such as metabolic or oxygen restrictions, or the filter-feeding efficiency of the lophophore. A research line starting-point is thus herein introduced which can lead to shed light into the adaptive strategies of rhynchonellides, relating particular morphotypes with potential triggering factors of mass extinction events.
... 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). ...
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.
... The genus Aphelesia is represented mainly by Aphelesia bipartita, displaying a high morphological variability (Fig. 4). Most of the individuals show the typical strongly biconvex profile and the distinctive subpentagonal to rounded-triangular outline illustrated by previous authors (Brocchi 1814;Davidson 1870;Cooper 1959;Gaetani and Saccà 1985a, b;Bitner and Martinell 2001;Bitner and Moissette 2003;Bitner and Dulai 2004), also showing a large and suberect beak and hypothyrid pedicle foramen. The low and trapezoidal uniplication in the anterior margin is distinctive of this species. ...
... Moreover, the ecological conditions prevailing in the basins where this assemblage was previously recorded consist of shallow-water, oxygenated, and high-energy bottoms, which range from circalittoral habitats (Gaetani and Saccá 1985a, b;Gaetani 1986;Bitner and Martinell 2001;Bitner and Moissette 2003), from subtidal, even subtidal peri-reefal, environments (Llompart and Calzada 1982) to the upper bathyal zone (Taddei Ruggiero 1994). In the Mediterranean-Atlantic seaway, Toscano-Grande et al. (2010) also pointed out that the depositional environment of these Tortonian-Pliocene brachiopod-bearing sandy deposits consists of shallow-water, high-energy habitats, exposed to wave action and currents, with sporadic storm events. ...
... Borghi (2001) also identified a Terebratula ampulla-Aphelesia bipartita assemblage from the Pliocene of Emilia (Italy). According to Bitner and Moissette (2003) the assemblages dominated by A. bipartita, T. terebratula and M. eusticta are characteristic of shallow-water, high-energy environment and sandy facies in Italy, Spain and North Africa (Gaetani and Saccá 1985a, b;Gaetani 1986;Taddei Ruggiero 1996;Bitner and Martinell 2001;Bitner and Moissette 2003). ...
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The paleogeography of the Late Neogene Atlantic-Mediterranean seaway via the Betic-Rifean Domain is quite complex due to the presence of several marine corridors. The study of transitional basins in these seaways is crucial to understand the configuration and evolution of the Mediterranean-Atlantic inter-connection. A mixed skeletal-siliciclastic sandstone succession located in one of these transitional areas (Guadix Basin, Southern Spain) was studied from a comprehensive paleontological standpoint focused on the main benthic assemblages (foraminifera, brachiopods, and trace fossils), integrating the data with the study of planktic foraminifera for an accurate biostratigraphic framework. Brachiopods are mostly represented by the Aphelesia-Gryphus assemblage. Two trace fossil assemblages were observed, dominated by Ophiomorpha with Bichordites (1) and Macaronichnus (2), respectively. The benthic foraminiferal assemblage is mostly represented by Planulina and Cibicides. The data gathered from the benthic communities reveal habitats with high-energy and turbulent conditions in an outer neritic-upper bathyal bathymetric range. Brachiopods from the Alicún section show a Mediterranean paleobiogeographic affinity. They were constrained in the Late Tortonian to the restricted basins of the Betic-Rifean Seaway and after the Messinian Salinity Crisis proliferated in both Mediterranean- and Atlantic-type basins of the Betic-Rifean Domain. The Guadix Basin contributed to the Mediterranean-Atlantic faunal inter-connection through the Betic-Rifean Seaway during the Late Tortonian and facilitated the earliest Pliocene expansion of brachiopods in the Mediterranean.
... Also the documented micromorphic brachiopods indicate post-mortem transport. Fossil Megerlia preferred shallow waters in high-energy environments (Bitner and Martinell 2001 and references therein;Bitner and Moissette 2003), similar to Agyrotheca which settled in cryptic habitats of firm substrates of the infra-to circalittoral (Dulai 2007). ...
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The Leitha Mountains in Austria are a chain of hills separating the southern Vienna Basin and the Danube Basin. The Lower East Alpine basement of the Leitha Mts. is covered by Middle Miocene sediments of the Badenian and Sarmatian regional stages. Close to the northeastern margin of these hills, upper Badenian successions are exposed, which are part of a coralline algal-dominated carbonate platform with hydrodynamically influenced sediments. Six sections have been logged and subjected to detailed investigation and sampling. They are characterized by inclined beds (foresets), which have been formed by unidirectional transport of sediments. Large-scale asymmetrical ripples indicate strong currents affecting shallow topset deposits. Generally, this hydrodynamically controlled sedimentation, documented by seven facies types, is reflected in a strongly reduced diversity of facies and biota, contrary to the older facies-rich middle Badenian sediments. This change from biologically to hydrodynamically controlled sedimentation led to a reduction in diversity of facies and biota. Sediment transport, however, caused secondary mass occurrences of echinoids or foraminifers derived from seagrass meadows. This study unravels the distribution and differences of middle and upper Badenian deposits of the Leitha Mountains and the influence of tectonic activity. Changes in hydrodynamics on the Leitha Platform are linked to the formation of the Danube Basin starting in the middle Badenian when a new seaway to the southeast has started to form.
... We evaluate depositional and taphonomic conditions that led to the origin of conspicuous, meter-scale shell concentrations formed by the large-sized brachiopod species Terebratula terebratula in a well-documented Upper Tortonian (Upper Miocene) paleogeographic seaway connecting the Atlantic Ocean and the Mediterranean Sea (Lee et al. 2001;García-Ramos 2006;Toscano-Grande et al. 2010). This species was very widespread and common in the Mediterranean region in the Miocene and Pliocene (Bitner and Moisette 2003), and many other species of the genus Terebratula occupied diverse environments and frequently formed a dominant Fig. 1 a, b Geographic location of the Guadix Basin in the Betic Cordillera (southern Spain). c Detailed map of the study area in the northern part of the Guadix Basin. ...
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Carbonate production by brachiopods in shallow-water habitats is generally expected to be not sufficiently high and temporally persistent to allow them to form very thick and densely packed shell concentrations. The formation of thick brachiopod concentrations requires long-term persistence of populations with high density of individuals, and such circumstances are assumed to be rare especially during the Cenozoic. However, here we show that the large-sized brachiopod Terebratula terebratula, the most common species in benthic assemblages with epifaunal bivalves and irregular echinoids, formed several decameter- to meter-thick, densely packed concentrations in shallow siliciclastic, high-energy environments, in a seaway connecting the Atlantic Ocean with the Mediterranean Sea during the Latest Tortonian (Late Miocene, Guadix Basin, southern Spain). This brachiopod formed (1) meter-scale, thick, parautochthonous concentrations in a prodelta setting and (2) thin, mainly allochthonous, tide- and storm-reworked concentrations in megaripples and dunes. The abundance of brachiopods at the spatial scale of the Guadix Basin seems to be mainly related to intermediate levels of sedimentation rate and current velocity because abundance and thickness of shell concentrations decline both (1) in onshore direction towards delta foresets with high sedimentation rate generated by debris flows and (2) in offshore direction with increasing levels of tide- and storm-induced substrate instability. Although brachiopods in dune and megaripple deposits are more fragmented, disarticulated, and sorted, and have a higher pedicle/brachial valve ratio than in prodelta deposits, taphonomic damage is still relatively high in prodelta deposits. Terebratula terebratula thus formed thick concentrations in spite of that disintegration processes were relatively intense along the whole depositional gradient. Therefore, population dynamic of this species was probably characterized by production maxima that were comparable to some Cenozoic molluscs in terms of their productivity potential to form thick shell concentrations in shallow subtidal environments. We suggest that temporal changes in brachiopod carbonate production have a significant spatial and phylogenetic component because multiple large-sized species of the family Terebratulidae, which underwent radiation during the Cenozoic, attained high abundances and formed shell concentrations in temperate regions.
... En las últimas décadas, numerosos estudios han puesto de manifiesto las importantes aportaciones de los braquiópodos al conocimiento de la paleoecología y bioestratigrafía de los sedimentos neógenos del sur de Europa y norte de África. Estas investigaciones han permitido delimitar las afinidades paleobiogeográficas de las faunas (Bitner y Moissette, 2003 ), efectuar reconstrucciones paleoambientales (Rico-García, 2007), realizar detallados análisis tafonómicos y de tafofacies (Rico-García et al., 2008) o desarrollar modelos de cálculo de edades en función de las velocidades de racemización de aminoácidos de sus conchas (Torres et al., 2000). La depresión del Guadalquivir es una de las cuencas neógenas más importantes de España, con una situación privilegiada para inferir la evolución paleogeográfica de las conexiones entre el océano Atlántico y el mar Mediterráneo. ...
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This paper analyzes the first brachiopod record of three Neogene formations of the southwestern Guadalquivir Basin (Huelva province, SW Spain) with a wide cartographical distribution. Samples obtained in five sections of these formations have allowed to recognize three species belonging to the genera Terebratula, Maltaia and Cryptopora, the latter being the first record of this genus in Spain. In addition, the biostratigraphical distribution and palaeoecological context of these species and others present in upper Neogene sediments (Tortonian-Lower Pliocene) of south Spain are discussed. Se analizan los primeros registros de braquiópodos de las tres formaciones geológicas de mayor distribución cartográfica en el suroeste de la cuenca del Guadalquivir (provincia de Huelva, sur de España). El muestreo realizado en cinco secciones ha permitido reconocer tres especies pertenecientes a los géneros Terebratula, Maltaia y Cryptopora, siendo este último el primer registro del género en España. Se discute el rango bioestratigráfico y el contexto paleoecológico de estas especies y otras halladas en sedimentos del Neógeno superior (Tortoniense-Plioceno Inferior) del sur de España.
... The outermost primary layer is of a thin granular nature, followed by a thin fibrous secondary and a thick prismatic tertiary layer (Fig. 1). In peripheral areas of the shell the tertiary layer completely pinches out and the secondary layer becomes dominant (Williams 1968;MacKinnon and Williams, 1974;Benigni 1985;Gaspard 1986;Bitner and Moissette, 2003). Gryphus vitreus lives from the shelf break at 150 m to about 300 m depth (Emig 1987(Emig , 1989a. ...
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Brachiopod shells are widely used as an archive to reconstruct elemental and isotopic composition of seawater. Studies, focused on oxygen and carbon isotopes over the last decades, are increasingly extending to the emerging calcium isotope system. To date, only little attention has been paid to test the reliability of fossil brachiopods on their modern counterparts.In this context, the present study investigates two modern brachiopods, Terebratulina septentrionalis (eastern Canada, 5–30 m depth, 7.1 °C seasonal temperature variation, two-layer shell) and Gryphus vitreus (northern Mediterranean, 200 m depth, constant all-year round temperature, three-layer shell). Both species were sampled along the ontogenetic growth direction and calcium, oxygen, and carbon isotopes as well as elemental concentration were measured. Calcium isotopes were analyzed on TIMS. The elemental composition was analyzed by LA-ICP-MS and ICP-AES.The results indicate an intra-specimen δ <sup>44/40</sup>Ca variation ranging from 0.16 to 0.33‰, pointing to a fairly homogenous distribution of calcium isotopes in brachiopod shells. However, in the light of the suggested 0.7‰ increase in calcium isotopes over the Phanerozoic such intra-specimen variations constrain ocean reconstruction. δ <sup>44/40</sup>Ca values of T. septentrionalis do not seem to be affected by growth rate. Calcium isotopic values of G. vitreus are heavy in the central part of the shell and trend towards lighter values in peripheral areas approaching the maximum isotopic composition of T. septentrionalis . The maximum inter-species δ <sup>44/40</sup>Ca difference of 0.62‰ between T. septentrionalis and G. vitreus indicates that care should be taken when using different taxa, species with different strontium content or brachiopods with specialized shell structure, such as G. vitreus , for ocean water reconstruction in terms of Ca isotopic composition. T. septentrionalis may record Ca isotopic fractionation related to seasonal seawater temperature variations in its shell but this is difficult to resolve at the current analytical precision. Average δ ¹⁸O-derived temperatures of the two investigated species are close to on-site measured temperatures.
... Fossil Megerlia are concentrated around the Mediterranean (Figs 3, 4), in southern and central Europe and North Africa (Logan et al. 2004). M. eustica has been recorded from Italy, Spain, and northwest Africa, and is apparently restricted to the Pliocene (Bitner & Martinell 2001;Bitner & Moissette 2003). The species is said to be characteristic of high energy, shallow water environments. ...
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A review is presented of the genera and species, both living and fossil, comprising the Superfamily Kraussinoidea and of their geographic and stratigraphic distributions. Two new fossil species are described, Kraussina chilensis and Megerlina miracula, from the Miocene of Chile and New Zealand respectively. The ontogenetic development of the brachidium in each of the four kraussinid genera is described and on the basis of recognised differences it is proposed that a new taxonomic subdivision is made that places Megerlia in a new subfamily Megerliinae, distinct from the other three genera in the existing subfamily Kraussininae Dall. The loop ontogeny of kraussinoids is compared with that of other terebratellidines and a phylogenetic linkage between kraussinoids and laqueoids is suggested.
... Occurrence. The genus Gryphus is recorded from the Pliocene and Pleistocene of Italy, Sacca 1984,1985), of the Pliocene of Spain (Bitner and Martinell2001) and the Pliocene of north-western Africa (Bitner and Moissette 2003). ...
... It is very common in the Miocene of the Paratethys (Bitner 1990) and the Pliocene of Italy Sacca 1985, Taddei Ruggiero 1994) and it was described from the Pliocene of North Africa (Bitner and Moissette 2003). In RJlOdes, it has been also mentioned from the Pliocene and described from the Pleistocene deposits by Spanopoulou (1997 Mmphology. ...
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Six brachiopod species (5 terebratulids and 1 craniid) from the upper Pliocene sandy and calcareous deposits of the Cape Vagia (Rhodes Island) are described: Gryphus sp., Terebratulina refusa, Megerlia truncata, Megerlia truncata var. monstruosa, Argyrotheca cordata, Megathiris detruncata and Novocrania anomala. This is the first record of Megerlia truncata var. monstruosa from the Pliocene of Greece and of Megerlia truncata from the Pliocene of Rhodes. Megathiris detruncata, Terebratulina retusa and Argyrotheca cordata are described for the first time from the Pliocene of Rhodes. This brachiopod association displays close affinity with those from the Mediterranean province, where assemblages of similar composition are associated with shallow-water, high energy environments. However, the association of Gryphus sp., Terebratulina retusa and Novocrania anomala corresponds to deeper water moderate environments.