Middle Triassic carbonate platforms in eastern Iberia: Evolution of their fauna and palaeogeographic significance in the western Tethys

Palaeogeography, Palaeoclimatology, Palaeoecology 417 (2015) 236–260

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Middle Triassic carbonate platforms in eastern Iberia: Evolution of their fauna and palaeogeographic significance in the western Tethys M.J. Escudero-Mozo a,b,⁎, A. Márquez-Aliaga c, A. Goy b,d, J. Martín-Chivelet a,b, J. López-Gómez b, L. Márquez c, A. Arche b, P. Plasencia c, C. Pla c, M. Marzo e, D. Sánchez-Fernández a a

Departamento de Estratigrafía, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain Instituto de Geociencias (UCM, CSIC), c/ José Antonio Nováis 12, 28040 Madrid, Spain Departamento de Geología, Facultad de Biología and Instituto Cavanilles, Universidad de Valencia, c/ Dr. Moliner 50, 46100 Burjassot, Valencia, Spain d Departamento de Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain e Department d'Estratigrafia, Paleontologia i Geosciènces Marines, Facultad de Geología, Universitat de Barcelona, 08008 Barcelona, Spain b c

a r t i c l e

i n f o

Article history: Received 8 May 2014 Received in revised form 14 October 2014 Accepted 31 October 2014 Available online 8 November 2014 Keywords: Ammonoids Bivalves Foraminifera Conodonts Anisian Ladinian

a b s t r a c t This article reports the first integrated study of the Middle Triassic of Iberia, based on the stratigraphy, sedimentology, and fossil fauna of Muschelkalk facies of the Iberian Ranges and the Catalan Coastal Ranges in Spain. On the basis of this study, new palaeogeographic reconstructions of the westernmost Tethys are proposed, and the evolution of the different palaeogeographic domains of Iberia (e.g., Iberian, Mediterranean, and Levantine– Balearic) are described. In these domains, Muschelkalk facies record the development of wide carbonate platforms that were the consequence of the first two broad marine transgressions of the Mesozoic in Iberia, respectively, late Pelsonian–early Illyrian and late Illyrian–Longobardian. Of these marine incursions, the oldest only manifested in the Mediterranean Triassic domain (Catalan Coastal Ranges and part of the Iberian Ranges), which acted as a palaeogeographic gulf opening northwards. Most of the fauna related to this first incursion show strong affinities with the Alpine/ Germanic bioprovinces, related to the Palaeotethys. In contrast, the second transgressive episode took place in a new regional palaeogeographic setting related to the intra-Pangea dextral shear, and the northward movement of the Cimmerian microcontinent. A rapid sea-level rise induced generalised marine flooding of the Iberian, Mediterranean, and Levantine–Balearic Triassic domains. The resulting carbonate platforms yield fossil assemblages (ammonoids, bivalves, foraminifera and conodonts) that show affinities with those of both the Alpine and Sephardic bioprovinces related to the Neotethys. These assemblages point to a significant increase in diversity during the late Fassanian–Longobardian, possibly related to the prevailing wider connections between the sea corridors, an increased continental run-off and input of nutrients and/or a general cooling of marine waters. © 2014 Elsevier B.V. All rights reserved.

1. Introduction The Middle Triassic was an exceptional time period during which, after the end-Permian mass extinction, several global factors determined an acceleration in the recovery of the biota and extensive radiation (Erwin, 1996). This key process in the evolution of life, which ended with the replacement of the “Palaeozoic Fauna” with the “Modern Fauna” (Sepkopski, 1984; Márquez-Aliaga, 2010; Ros et al., 2011), was triggered by the progressive stabilisation of the carbon cycle, which had been punctuated during the Early Triassic by multiple periods of massive CO2 release (Payne and Kump, 2007). This strong

⁎ Corresponding author at: Departamento de Estratigrafía, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain. E-mail address: [email protected] (M.J. Escudero-Mozo).

http://dx.doi.org/10.1016/j.palaeo.2014.10.041 0031-0182/© 2014 Elsevier B.V. All rights reserved.

instability in the carbon cycle and the associated volcanic input of sulphurous gasses, provoked huge environmental perturbations in both the atmosphere and the oceans (Márquez-Aliaga et al., 2003; Kidder and Worsley, 2004; Márquez, 2005; Plasencia and MárquezAliaga, 2005; Woods, 2005; Tong et al., 2007; Sun et al., 2012) and hampered biotic recovery in immature and poorly functioning marine ecosystems throughout the Early Triassic. The delay in recovery lasted about 5–10 m.y. (Lehrmann et al., 2006; Pruss et al., 2006; Algeo et al., 2011), though some groups such as ceratitids and ammonoids, possibly recovered faster than was previously estimated (Brayard et al., 2009). At the time of environmental stabilisation, the western Tethys realm experienced a broad marine transgression from the East, controlled by the break-up of Pangaea. That marine invasion took millions of years to reach its westernmost boundary in Iberia. This occurred in the Middle Anisian (López-Gómez et al., 1998), when shallow marine waters

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determined the deposition of shallow marine carbonate units (Muschelkalk facies). Such marine carbonate deposition prevailed until the end of the Ladinian (or onset of the Carnian), when a prolonged regressive episode allowed for the deposition of the Keuper facies (López-Gómez et al., 2002). The present integrated study of the Middle Triassic carbonate ramps of E Iberia (Iberian Ranges and Catalan Coastal Ranges) (Fig. 1) based on stratigraphic, sedimentologic, and palaeontologic data sets, proposes a new stratigraphic framework that differs substantially from existing palaeogeographic reconstructions of the westernmost Tethys for the Middle Triassic. Marine fossil assemblages (ammonoids, bivalves, foraminifera and conodonts) were examined in terms of biogeographic affinities and changes in palaeodiversity. The results reported here should provide new insight into how life recovered in the shallow waters of the Tethys after the Permian mass extinction.

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2. Geological and geographical framework The evolution of the Triassic grabens systems of Western and Central Europe was related to the southward propagation of the NorwegianGreenland rift system and coeval with the development of the Tethys rift system (Ziegler, 1990), which took place during fragmentation of the Pangaea supercontinent. In this period of crustal extension, four major rift basins developed in the northern, northeastern and eastern portions of Iberia (Pyrenean, Ebro, Catalan and Iberian Basins), primarily through reactivation of older Variscan faults. The Iberian and Catalan Basins developed primarily during the Mesozoic, when they experienced several extensional periods (Arche and López-Gómez, 1996; De Vicente et al., 2009). The first of these periods (Late Permian–Late Jurassic) occurred in two main stages: a first episode or rifting stage with at least three synrift–postrift pulses characterised by

Fig. 1. Permian–Triassic outcrops in the Iberian Ranges and Catalan Coastal Ranges. Indicated are the geographical locations of the main fossiliferous sections and palaeontological sites of the Muschelkalk facies. Stratigraphic sections of the Iberian Ranges: 1. Huélamo; 2. Valdemeca; 3. Camarena; 4. Barranco de la Hoya; 5. Cerro Morrón; 6. Boniches; 7. Henarejos; 8. Villora; 9. La Ermita; 10. Moya; 11. El Paraíso; 12. Chelva; 13. El Molinar; 14. Cueva Cirat; 15. El Tormo; 16. Serra; 17. Agujas de Santa Águeda; 18. Jarafuel; 19. Mijares; 20. Macastre; 21. Montserrat; 22. Bugarra; 23. Calanda. Main palaeontological sites of the Catalan Coastal Ranges: 24. Alfara; 25. Benifallet; 26. Camposines; 27. Begues; 28. Olesa; 29. El Farrel, 30. Centelles and 31. L'Ametlla.

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continental deposits mainly of alluvial origin from Middle–Late Permian to Early Triassic, and a second thermal subsidence episode with many synrift–postrift pulses extending (Vargas et al., 2009) until the Late Jurassic. This latter stage took place during transgression of the Tethys Sea in the eastern Iberian basins and deposition of marine Muschelkalk facies during the Middle Triassic. These facies are examined in this paper. The Triassic stratigraphic record in the Iberian and Catalan Basins is broadly represented by the Buntsandstein, Muschelkalk and Keuper facies (typical Germanic Triassic). Muschelkalk facies in the central part of the Iberian Basin are represented by two marine carbonate units: the Landete Formation and the Cañete Formation, which are separated by a unit of mixed evaporite–siliciclastic composition (Mas Formation) (López-Gómez et al., 1993) (Fig. 2). Three similar units were described in the Catalan Basin: the “Lower”, “Middle”, and “Upper” Muschelkalk (Calvet and Ramón, 1987; Calvet and Marzo, 1994) (Fig. 2). This lithologic “trilogy” has been traditionally called the “Mediterranean Triassic” (Virgili, 1977). In the westernmost part of the Iberian Basin (close to the Hesperian massif), only one marine carbonate level is found. This level, or Cañete Formation, rests conformably or unconformably on different Early Triassic continental units or on the Variscan basement. The succession was named “Iberian Triassic” by Virgili (1977) to differentiate it from the Mediterranean Triassic (Fig. 2). Interestingly, in the easternmost and southernmost parts of the Iberian Ranges, Muschelkalk facies succession is also represented by a single carbonate unit, a singular feature that led López-Gómez et al. (1998) to propose a new palaeogeographic domain characterised by the “Levantine–Balearic Triassic”. In that proposal, the presence of just one carbonate unit instead of the two carbonate units of the “Mediterranean Triassic” plus the intermediate siliciclastic–evaporite unit was interpreted

as the result of regional disappearance, due to a lateral facies change into marine carbonates, of the siliciclastic–evaporite unit, and the consequent amalgamation of the under and overlying carbonate bodies. Herein, a new stratigraphic scheme is proposed for the Levantine– Balearic Triassic, based on new palaeontological and stratigraphic data sets. In this scheme, the single carbonate level of the Levantine–Balearic Triassic is interpreted to be stratigraphically equivalent to exclusively its upper carbonate level (the Cañete Formation or Upper Muschelkalk unit) (Fig. 2). This new stratigraphic interpretation has important implications for palaeogeographic reconstructions of Iberia and the western Tethys. Based on fossil contents at the top of the Cañete Formation, this domain can be divided into: a northern part characterised by a low fossil content, and a southern part characterised by a high fossil content. In addition, the Catalan Basin has been divided into three palaeogeographic sub-domains limited by NW-SE fault systems: from SW to NE, the Priorat–Baix Ebre, Prades and Gaiá–Montseny domains (Marzo, 1980; Marzo and Calvet, 1985) (Fig. 1). These Middle Triassic units (Muschelkalk facies), combined with the uppermost part of the Buntsandstein (or Röt) facies and the lower part of the Keuper facies, record two major transgressive–regressive cycles (TR-1 and TR-2) (Calvet et al., 1990; López-Gómez et al., 1993) in both the Iberian and Catalan Basins (Fig. 2). The first one (middle–late Anisian) includes the Röt facies, the Landete Formation and the lower part of the Mas Formation in the Iberian Basin; the “Lutite, Carbonate, Evaporate unit” (Röt facies), the Lower Muschelkalk and part of the Middle Muschelkalk in the Catalan Basin (Fig. 2). The second cycle (latest Anisian and Ladinian) recorded throughout the study area comprises the upper part of the Mas Formation, the Cañete Formation and the lower part of the Keuper in the Iberian Basin, as well as the upper part of the Middle Muschelkalk, the Upper Muschelkalk and the first levels of the Keuper in the Catalan Basin (Fig. 2).

Fig. 2. Chronostratigraphic chart of the Triassic formations of the Iberian and Catalan Coastal Ranges and depositional sequences. T–R.1: First transgressive–regressive cycle; T–R.2: Second transgressive–regressive cycle; VLD: Valdemeca unit; AC: Cañizar Sandstones Formation.; LAE: Eslida Siltstones and Sandstones Formation; CPS: Upper Prades Conglomerates unit; API; Lower Prades Sandstones unit; APS: Upper Prades Sandstones unit; CGS; Upper Garraf Conglomerates unit; AE: L'Eramprunyá Sandstones unit; ALA: L'Aragall Sandstones and Shales unit; ACC: Caldes Conglomeratic Sandstones unit; ALF: Figaró Sandstones and Shales unit; CLCES; “Lutite, Carbonate, Evaporite” unit. Data for the continental Lower Triassic units are based on Calvet and Marzo (1994), Galán-Abellán et al. (2013) and López-Gómez et al. (2012).

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After Alpine compressional tectonics in Iberia during the Cenozoic, the Iberian Basin gave rise to the present-day Iberian Ranges (IR) and the Catalan basin to the Catalan Coastal Ranges (CCR). The IR can be geographically divided into two sectors or “branches”: the Castilian Branch to the SW and the Aragonese Branch to the NE with the Cenozoic depression of Almazan–Teruel in between (López-Gómez et al., 2002). The Iberian Ranges are controlled by two major fault systems: a longitudinal one trending NW-SE (Variscan, reactivated during alpine tectonics) and a transverse one trending NE-SW (Alpine), which developed mainly in the eastern part of the ranges (Arche and López-Gómez, 1996) (Fig. 1). The CCR are a Cenozoic NE-SW structure dominated by longitudinal, near vertical basement faults with a NE-SW to ENE-WSW trend; some of these structures are deviated by a set of strike-slip faults trending NW-SE (Anadón et al., 1979; Guimerá, 1984, 1988; Gaspar-Escribano et al., 2004). 3. Material and methods For this study, bibliographic and palaeontological data were reviewed in detail and compiled from the Muschelkalk facies in the Iberian and Catalan Coastal Ranges. The stratigraphic sections reviewed were 23 in the IR, one in the Aragonese Branch (Calanda) and 22 in the Castillian Branch (Fig. 1). In the latter branch, the sections belong to different domains (Fig. 1) and consist of: A — twelve sections from the Mediterranean Triassic domain, including seven from the Landete Formation and five from the Cañete Formation; B — five sections in the northern part of Levantine–Balearic Triassic domain; and C — five sections in the southern part of the Levantine–Balearic Triassic domain. In each of these domains, a synthetic section was constructed for each area based on a stratigraphic study and correlations with the sections showing the precise stratigraphic positions of the fossil horizons. Although the Calanda section forms part of the Iberian Ranges, in some sections of this work, it is considered independent of this geographical area due to its palaeogeographic position (intermediate between the Iberian and Catalan Basins) and its high fossil content. In the Catalan Coastal Ranges, one stratigraphic section (Venta de Camposines) was surveyed for this study and eight palaeontological sites previously studied by Virgili (1958) and Márquez-Aliaga (1985) were revised (Fig. 1). Three of these sites (Alfara, Benifallet and Camposines) occur in the Baix Ebre domain and correspond to the Upper Muschelkalk unit. The other five (Begues, Olesa, El Farrel, Centelles and L'Ametlla) belong to the Gaiá–Montseny domain and correspond to the Lower Muschelkalk unit (Fig. 1). Most of the palaeontological material of this area comes from museum collections, and the precise stratigraphic positions of their fossils are usually

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unknown; only the members of the Lower or Upper Muschelkalk in which they were found are referenced. Using this information along with detailed stratigraphical and sedimentologic studies, Calvet et al. (1990) and López-Gómez et al. (1998) built a synthetic stratigraphical section in which the palaeontological material was allocated to the most accurate position possible. Based on the information obtained from these general data, specimens and palaeocommunities are here described. Results are established for the different domains, as well as through comparisons between these domains and others in the western Tethys area. 4. Lithostratigraphy The lithostratigraphy for the Iberian and Catalan Coastal Ranges is based on the units of both areas from the oldest to the youngest unit in each case. Some of the main stratigraphic sections studied were then used to build the synthetic sections of each domain of the Iberian Ranges shown in Figs. 3 and 4. 4.1. Iberian Ranges The Landete Formation is the lower carbonate unit of the IR. Slight variations in thickness are seen across the study area (30 to 49 m), decreasing towards the NW in the Valdemeca area where the Landete Formation pinches out (Iberian Triassic domain) (Figs. 1, 2). The age of the unit is Pelsonian–Illyrian (middle–late Anisian) (López-Gómez et al., 1998), and it only crops out in the Mediterranean Triassic domain, where it overlies different continental units, unconformably (Cañizar Formation) or conformably (Röt facies) (Fig. 2). The Landete Formation primarily consists of decimetre-scale banks of well-bedded grey dolomicrites with sandy dolomite banks at the base of the unit. Occasionally, this unit contains intercalations of small horizons of grey–yellow clay dolomites. Most of the unit is dolomitised, hiding the original textures and lithologies of the rocks. However, original characteristics are occasionally preserved, including remains of foraminifers and bivalves. This formation was the result of shallow marine carbonate deposition in a ramp setting, ranging from subtidal (typically of high-energy, with generation of shoals) to supratidal sabkhas (López-Gómez et al., 1993, 1998). The formation is divided into six members based on sedimentological characteristics and palaeogeographic distributions. From base to top, these subunits are as follows: 1. Serra Member (peritidal facies); 2. San Martín Member (high-energy subtidal to intertidal); 3. Mal Paso Member (low energy subtidal to intertidal); 4. Olocau Member (upper subtidal–intertidal to intertidal and sporadically lower

Fig. 3. Some of the major stratigraphic sections of the Landete Formation used to build the synthetic section. Camarena, Barranco de la Hoya and Chelva sections modified from López-Gómez, (1985). For legend, see Figs. 4 and 5.

240 M.J. Escudero-Mozo et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 417 (2015) 236–260 Fig. 4. Some of the major fossiliferous stratigraphic sections of the Cañete Formation in the IR used to build the synthetic series of each domain. The Calanda section is modified from Márquez-Aliaga et al. (1994); the Agujas de Santa Águeda section is modified from López-Gómez et al. (2005); and the Bugarra and Montserrat sections are modified from Sánchez-Fernández et al. (2005). The fossil legend is shown in Fig. 5.

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supertidal); 5. Peña Rubia Member (upper intertidal to lower supratidal); and 6. Beamud Member (supratidal facies dominated by shabkha conditions) (López-Gómez and Arche, 1992). The Cañete Formation crops out along the Iberian Ranges, ranging in thickness from 37 to 130 m, and decreasing towards the northwest. The lower contact is transitional with the Mas Formation in the western area (Mediterranean Triassic domain) and most likely overlays the Buntsandtein facies (Röt facies) in the eastern area (Levantine–Balearic Triassic domain) (Fig. 2). Farther to the west, in the Cueva de Hierro, the Cañete Formation lies unconformably over the basement. The upper contact is always transitional to the Keuper facies (López-Gómez et al., 1993) (Fig. 2). This unit essentially consists of grey dolomite and limestone with smaller amounts of marls and green–yellow clay. Dolomites, which predominate at the base of the unit, show a fine-medium grain size and a grey–ochre colour. Although most primary features were blurred by dolomitization, oolitic and tractive levels with quartz grains can be occasionally recognised. Limestones are commonly fine grained, and include towards the top of the unit, several levels of high fossil contents, yielding bivalves, ammonoids, foraminifers, conodonts, and chondricthyans. Most of this unit has been dated as Ladinian by means of ammonoids and conodonts (López-Gómez et al., 1998). The Cañete Formation records the development of a large shallow carbonate ramp. The formation is divided into five members defined by their sedimentological characteristics and their palaeogeographic distribution. These are (from bottom to top): 1. Gorgocil Member (high-energy to intertidal facies); 2. Henarejos Member (open proximal platform to protected subtidal); 3. Huélamo Member (low-energy subtidal to intertidal); 4. Valacloche Member (tidal channels); and 5. Moya Member (shallow protected subtidal, intertidal and supratidal) (López-Gómez and Arche, 1992; López-Gómez et al., 1993). 4.2. Catalan Coastal Ranges The Lower Muschelkalk in the Catalan Coastal Ranges consists of limestone and dolomite, ranging in thickness from 70 to 120 m in the northeastern and southwestern areas, respectively. The unit shows a transitional lower contact with the Lutite, Carbonate, and Evaporite Unit (Röt facies), as well as a transitional upper contact with the clastic–evaporitic facies of the Middle Muschelkalk (Fig. 2). Based

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on ammonoids and conodonts, the unit is dated as Pelsonian–Illyrian (middle–late Anisian) (Calvet and Marzo, 1994; Goy, 1995; López-Gómez et al., 1998; Márquez-Aliaga et al., 2000). The unit arose from the shallow to open marine deposition of carbonates. The unit is divided into four members (Calvet and Ramón, 1987; Calvet et al., 1990; Calvet and Marzo, 1994) from base to top: 1. El Brull Member (shallow subtidal to supratidal deposits); 2. Olesa Member (lagoon bioclastic carbonates); 3. Vilella Baixa Member (open marine burrowed deposits); and 4. Colldejou Member (white peritidal dolomite facies). The first three members are grey in colour and may be totally or partially dolomitised (Calvet et al., 1990). The Upper Muschelkalk ranges in thickness from 100 to 140 m in the northwest and southwest, respectively. This unit has a transitional lower contact with the clastic–evaporitic facies of the Middle Muschelkalk unit and a transitional upper contact with the Keuper facies (Fig. 2). Several horizons of the unit are dated as Ladinian by ammonoids and conodonts (Calvet and Marzo, 1994; Goy, 1995). This unit has been divided into several members in each domain of the CCR (Calvet et al, 1987; Calvet et al., 1990; Calvet and Marzo, 1994) and new interpretations have been recently provided by MercedesMartín et al. (2013). In the Baix-Ebre domain, which is the only one examined here, the unit is divided from base to top as follows: 1. Rojals Member, (oolitic limestones and dolomites, subtidal to supratidal); 2. Benifallet Member (bioturbated limestones and dolomites, from subtidal environments); 3. Rasquera Member (open marine limestones, dolomites and shales with Daonella); 4. Tivissa Member (outer to shallow ramp carbonates); and 5. Capafons Member (dolomite, marls, shales, and breccias of peritidal settings). 5. Fossil contents The fossil contents of the Middle Triassic Muschelkalk facies of the Iberian Ranges and Catalan Coastal Ranges have been addressed by authors and research groups since the 19th century: De Verneuil and Collomb (1853), De Verneuil (1854), Mojsisovics (1882, 1887), Würm (1911, 1913), Tornquist (1916), Villaseca (1920), Bataller and Guerin (1930), Schmidt (1936), Bataller (1954), Virgili (1958), Márquez et al. (1994), Goy (1995), Márquez-Aliaga (1985), Márquez-Aliaga and Martínez (1996), Márquez (2005), Ros-Franch (2009) and Plasencia (2009).

Fig. 5. A. Synthetic section of the Landete Formation, Mediterranean Triassic domain in the Iberian Ranges showing the stratigraphic positions of the most representative fossils. B. Synthetic section of the Lower Muschelkalk of the Catalan Coastal Ranges modified from Calvet et al. (1990) showing the approximate stratigraphic positions of some of the most representative fossils.

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5.1. Ammonoids Only a few Anisian and Ladinian ammonoids from Iberia have been described and figured, mainly because of the predominance of shallow, often restricted, water facies and also because of the strong dolomitization that affects most of the units. For the Anisian, the IR have not yet yielded any ammonoids (Landete Formation). In the CCR, they have been found exclusively in the upper part of the Olesa Member (Lower Muschelkalk), where several specimens of Paraceratites and a few Olesites (Fig. 5B) have been found. These have been identified as Paraceratites occidentalis (Tornquist), P. flexuosiformis (Tornquist), P. evoluto-spinosus (Tornquist), P. catalanicus (Bataller), P. guerini (Bataller), P. almeari (Bataller) and Olesites villaltai (Virgili) (Virgili, 1958; Goy, 1995). This assemblage can be ascribed to a late Pelsonian (middle Anisian) and/or early Illyrian (late Anisian) age. Ladinian ammonoids are more abundant, and come from the Mediterranean Triassic domain. In the IR, most of the ammonoids have been recovered from several levels of the upper part of the Cañete Formation. According to their age and stratigraphic position, two main assemblages can be differentiated. The lower one is late Fassanian and contains some generalist specimens such as Eoprotrachyceras (E. cf. vilanovai

(D'Archiac)) in the Albarracín area and Gevanites archei (Goy) in the Henarejos (Fig. 11: 7), Moya and El Paraíso sections. The next higher assemblage is Longobardian and contains several taxa of the “Cephalopod Spanish fauna” from Mojsisovics (1882), including Protrachyceras hispanicum (Mojsisovics) and Iberites pradoi (D'Archiac) (Fig. 11: 8), together with Anolcites sp. and Iberites sp. (Fig. 6). The most complete ammonoid succession of the Ladinan in Iberia occurs in the Camposines area (the Baix-Ebre domain of the CCR, Fig. 7). This succession begins in the lower part of the Rasquera Member with the appearance of Eoprotrachyceras curionii (Mojsisovics), which marks the beginning of the Fassanian (E. curionii Biozone). A few metres above, the following assemblage is found: Eoprotrachyceras vilanovai (D'Archiac) (Fig. 11: 1), Anolcites ibericus (Mojsisovics), Gymnites cf. incultus Beyrich and Iberites sp., which indicates a late Fassanian age (E. vilanovai Biozone). Finally, several specimens of P. hispanicum (Mojsisovics) (Fig. 11: 2) associated with I. pradoi (D'Archiac) (Fig. 11: 5, 6) (Goy, 1986, 1995), which also appear together in the Alcover Member in the Prades domain, are at the limit of the Rasquera and Tivissa Members. P. hispanicum characterises the P. hispanicum Biozone and marks the onset of the Longobardian. However, some ammonoids cited as Protrachyceras in this area might correspond to species of the Anolcites genus, which are

Fig. 6. Synthetic section of the Cañete Formation in the Mediterranean Triassic domain of the Iberian Ranges showing the stratigraphic positions of the most representative fossils. BRA: brachiopods; CO: conodonts; FORA: foraminifers; GASTR: gastropods; CHONDRIC: chondrichthyan; IC: icnites.

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Fig. 7. Synthetic section of the Upper Muschelkalk of the Catalan Coastal Ranges (Baix–Ebre domain) modified from Calvet et al. (1990) showing the approximate stratigraphic positions of the most representative fossils.

also Longobardian (Goy, 1995). Virgili (1958) reported some Nannites (N. bittneri Mojsisovics and N. cf. fugax Mojsisovics) in the “limestone with Daonellas” (i.e., Cobaltó Member) of the Gaiá–Montseny domain. In the Calanda area (Fig. 8), a succession comparable to that of the Camposines area exists, although more incomplete. In this section, E. vilanovai (D'Archiac), Nannites mambrini Schmidt and Iberites sp. from the Fassanian and Protrachyceras cf. hispanicum (Mojsisovics) and Anolcites sp. from the Longobardian, have been cited (Anadón and Albert, 1973; Márquez-Aliaga et al., 1994). Southwards, in the northern part of the Levantine–Balearic Triassic domain, the El Molinar section has yielded the oldest ammonite of the IR (Fig. 9) found to date. This single poorly-preserved specimen might be attributed to Schreyerites (aff. S. abichi) (López-Gómez et al., 1998), which is Illyrian. In the Ladinian of this area, only ammonoids of lower Fassanian age have been found. In particular, in the stratigraphic sections of Agujas de Santa Águeda (López-Gómez et al., 2005) and El Tormo (Fig. 9), Eoprotrachyceras (E. cf. curionii), frequent Proarcestes and Flexoptychites have been identified. Finally, in the southern part of the Levantine–Balearic Triassic domain, only two ammonoids have been reported, both of from the Bugarra section: Proarcestes (Fig. 11: 3), which is located near the

base of the section (early Fassanian), and Anolcites cf. doleriticus (Mojsisovics) (Fig. 11: 4) found in its upper part (late Fassanian–early Longobardian) (Fig. 10). It should be noted that almost all the specimens found in the study area are endemic forms, with the exception of the cosmopolitan species E. curiioni, and A. cf. doleriticus, which also appear in other parts of the Tethys realm (Mietto and Manfrin, 1995). At the genus level, most of the reported genera (Eoprotrachyceras, Protrachyceras, Anolcites…) indicate clear influence of the Tethys realm. However, the specimens belonging to the Gevanites genus (Hungaritidae) appearing in the upper part of the Cañete Formation in the Mediterranean Triassic domain of the IR (upper Fassanian), indicate some input of fauna from the Sephardic bioprovince (Hirsch, 1977; Parnes et al., 1985; Hirsch et al., 1987). Biostratigraphically, the Muschelkalk facies of E Iberia contain ammonoid assemblages that can be included in three biozones (each starting with the lower appearance of the index species) roughly correlated with the standard Zones of the Tethys (Balini et al., 2010) (Fig. 12): a) the Eoprotrachyceras curionii Biozone, characterised by the presence of Eoprotrachyceras curionii and early Fassanian; b) the Eoprotrachyceras vilanovai Biozone, characterised by the presence of E. vilanovai and late

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Fig. 8. Calanda synthetic section (top of the Cañete Formation) showing the stratigraphic positions of the most representative fossils. Modified from Márquez-Aliaga et al., 1994. BR: brachiopods; CONOD: conodonts; GAST: gastropods.

Fassanian; and c) the Protrachyceras hispanicum Biozone, characterised by the presence of P. hispanicum and early Longobardian. 5.2. Bivalves Bivalve associations of Anisian age have been found in the Iberian Ranges in the Landete Formation (Mediterranean Triassic domain) and also at the base of the Cañete Formation (Serra section, Levantine– Balearic Triassic domain), and in the Catalan Coastal Ranges, in the Lower Muschelkalk. These associations, listed in Figs. 5A–B and 9, are characterised by their low diversity and are often composed of the dominant cosmopolitan specimens from the Alpine/Germanic bioprovinces (Palaeotethys) related to shallow and/or restricted environments. The most characteristic bivalves of the Serra section are shown in Fig. 13. The species described in these assemblages are commonly represented in several areas of Europe and Israel. In the Negev area (Israel),

Lerman (1960) described Myophoria vulgaris (Schlotheim), Elegantina aff. intermedia (Schauroth), Buermesia posteroradiata Cox, Unionites fassanensis Wissman and Neoschizodus orbicularis Bronn. In the Tolbouhin section (Bulgaria), Pleuromya elongata Schlotheim and Myophoria orbicularis Bronn are described from Encheva (1969). In the Pelsonian of Belogradchik, Myophoria vulgaris Schlotheim, Hoernesia socialis (Schlotheim), Unionites fassaensis Wissman and Pleuromya sp. were reported by Márquez-Aliaga (1985). In the lower Muschelkalk of the Polish Lowland, Senkowiczowa (1985) detailed several cosmopolitan species marked by a wide Triassic range that were related to muddy bottoms, including M. vulgaris (Schlotheim), Neoschyzodus laevigatus (Goldfuss), N. orbicularis Bronn, H. socialis (Schlotheim), Pleuromya sp. and U. fassaensis Wissman. In the Muschelkalk of the Mecsek Mts (Hungary), Szente (1997) described Pleuromya elongata Schlotheim, H. socialis (Schlotheim), N. laevigatus (Goldfuss) and M. vulgaris (Schlotheim). In the lower Muschelkalk of Germany (lower

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Fig. 10. Synthetic section of the Cañete Formation in the southern part of the Levantine–Balearic Triassic domain of the Iberian Ranges showing the stratigraphic positions of the most representative fossils. AMMO: ammonites; BRA: brachiopods; CONO: conodonts; GASTRO: gastropods; A: algae.

Bithynian–upper Aegean to lower Illyrian), Klug et al. (2005) cited M. vulgaris (Schlotheim), Neoschizodus laevigatus (Goldfuss), N. orbicularis Bronn and Trigonodus sandberger Alberti. For the Ladinian, bivalve assemblages have been described from the Mediterranean Triassic and the southern part of the Levantine–Balearic Triassic domains of the IR. These assemblages, listed in Figs. 6 and 10, show a greater diversity than those from the Anisian. The bivalves yielded from the first fossiliferous levels of the Ladinian still show a relatively low diversity in all areas: only three species are reported from the Mediterranean Triassic domain (Leptochondria alberti, Pseudocorbula gregaria and Pseudoplacunopsis teruelensis) and five from the southern Levantine–Balearic Triassic domain (Bakevellia costata, Elegantina sublaevis, Limea vilasecai, Modiolus sp. and Pseudocorbula gregaria). These bivalves were pioneer taxa and probably lived under stressful environmental conditions. The brachiopod Lingularia, an opportunist form, is also common in some of these levels. The bivalve assemblages of the Ladinian (Cañete Formation), are richer in diversity and show biogeographic affinities with two bioprovinces of the Tethys realm. The assemblages include benthonic cosmopolitan Tethyan bivalves of Alpine affinity, such as Pseudocorbula gregaria (Muster), Modiolus myconchaeformis (Phillipp), Bakevellia costata (Schlotheim), Bakevellia subcostata (Goldfuss), Bakevelia crispata (Goldfuss), Unionites muensteri (Wissman), Neoschizodus laevigatus (Goldfuss), Leptochondria albertii (Goldfuss), Umbostrea

cristadifformis (Schlotheim), Entolium discites (Schlotheim), Paleonucula goldfussi (Alberti), “Mytilus” eduliformis (Schmidt) and Paleoneilo eliptica (Goldfuss), and benthic bivalves characteristic of the Sephardic bioprovince, which are also found in the Negev area (Israel), including Gervillia joleaudi (Schmidt), and Pseudoplacunopsis teruelensis Würm, as well as Elegantina sublaevis (Schmidt), Limea vilasecai (Schmidt) and Costatoria kiliani Schmidt (Hirsch, 1977; Márquez-Aliaga, 1985; Márquez-Aliaga and López Gómez, 1989; Márquez-Aliaga and Ros, 2003; Hirsch et al., 2014). Some of these bivalves are shown in Figs. 14 and 15. Assemblages consist of cosmopolitan epibentonic and some infaunal bivalves, characteristic of shallow and/or protected environments (lagoon). In the CCR, Ladinian bivalve assemblages contain cosmopolitan Tethyan bivalves of Alpine affinity (Fig. 7) including some species common to the IR. In this area, no species of Sephardic origin have been described. Based on their stratigraphic positions, it is possible to differentiated two associations: one in the Rasquera Member comprising B. subcostata, Chlamys sp., E. discites, Limea costata (Goldfussi), Elgantina sp. and P. elongate; and a second one in the Capafons Member that consists of Cassianela decussata (Munster), Costatoria goldfussi (Alberti) and E. sublaevis Schmidt. These Ladinian assemblages consist of cosmopolitan epibentonic and some infaunal bivalves characteristic of low energy subtidal environments (Rasquera Member) or shallow restricted environments (Capafons Member).

Fig. 9. Synthetic section of the Cañete Formation in the northern part of the Levantine–Balearic Triassic domain, Iberian Ranges showing the stratigraphic positions of the most representative fossils. B: brachiopods; F: foraminifers; P: porifers.

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Fig. 11. 1. Eoprotrachyceras vilanovai (D'Archiac), VC-6/1. Fassanian: E. vilanovai Biozone. Venta de Camposines, CCR. 2. Protrachyceras hispanicum (Mojsisovics), VC-18/4. Longobardian: P. hispanicum Biozone. Venta de Camposines, CCR. 3. Proarcestes sp., Bu-5/1. Fassanian: E. curionni Biozone. Bugarra, IR. 4. Anolcites cf. doleriticus (Mojsisovics), Bu-86/1. Longobardian: P. hispanicum Biozone. Bugarra, IR. 5, 6. Iberites pradoi (D'Archiac), VC-15/1,4. Longobardian: P. hispanicum Biozone. Venta de Camposines, CCR. 7. Gevanites archei Goy, He-18. Fassanian: E. vilanovai Biozone. Henarejos, IR. 8. Iberites pradoi (D'Archiac), Pa-25/1. Longobardian: P. hispanicum Biozone. El Paraíso, IR. All ammonites are shown as their actual sizes except specimen 4, which is enlarged ×1.5.

Some Alpine nektonic (Daonella lommeli (Wissman)) and planktonic (Bositra wengensis (Wissman)) elements have been also cited from the base of the Fassanian in the IR and the CCR (Budurov et al., 1993). Unlike the other assemblages, these species indicate open marine environments.

5.3. Foraminifers In the Anisian of the IR, very few foraminifers have been reported, mainly because of intense pervasive dolomitization of the original

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Fig. 12. Biochronostratigraphic chart showing the biostratigraphic distribution of ammonites in E Iberia, the Biozones proposed for this area, and equivalences with the Standard Tethys ammonoid Zones. Hu: Hungaritidae; G: Gymnitidae; A: Arcestidae; N.: Nannitidae; Ar.: Arpaditidae; Tr.: Trachyceratidae; P: Ptychitidae.

carbonate facies in the area. In particular, these have been described from the Landete Formation (Mediterranean Triassic domain) (Fig. 16: 1–4) and the basal levels of the Cañete Formation (in the Serra section, Fig. 16: 5) (Levantine–Balearic Triassic domain) (Márquez et al., 1994; Escudero-Mozo et al., 2012a). These foraminifers are listed in Figs. 5A and 9. In the Anisian of the CCR, however, several levels with foraminifers have been cited from the Olesa and the Vilella Baixa Members (Fig. 5B) (Budurov et al., 1993). Foraminifer assemblages consist of different species in each area and the only common form is Hoyenella sinensis (Ho). All of the assemblages are typical of the Alpine zone, and the fauna assemblages are comparable to those of the other regions of the Palaeotethys realm. The Anisian foraminifers of the IR are characteristic of protected lagoon environments and are considered part of the Pilammina densa Zone (in the sense of Salaj et al., 1988); the CCR assemblages are characteristic of the uppermost part of the same zone. Both the IR and CCR assemblages indicate a Pelsonian–Illyrian age (Pérez-Arlucea and Trifonova, 1993; Calvet and Marzo, 1994; Márquez, 2005). The Ladinian foraminifer assemblages have been described from the IR, CCR and Calanda section. In the Mediterranean Triassic domain (Henarejos section), only a single level containing H. sinensis (Ho) and Nodosaria ordinata Trifonova has been found (Fig. 6). In the southern part of the Levantine–Balearic Triassic domain of the IR, two high diversity assemblages, listed in Fig. 10, were reported at the base and top of the Cañete Formation by Sánchez-Fernández et al. (2005); some of which are shown in Fig. 16(6–10). Both these assemblages have some common species including abundant Involutinidae, which are frequent in the Alpine domain. These foraminifers point to low energy environments, except Planiinvoluta carinata Leischner, Tolypammina gregaria Wendt and Calcitornella sp., which are attached to different substrates and could indicate slightly higher energy environments (Márquez, 2005). A Longobardian–Carnian age was proposed by Trifonova (1993) for the Involutinidae species found in these assemblages Lameliconus ex gr. ventroplanus biconvexus (Oberhauser), Lamelliconus multiespirus (Oberhauser), Lameliconus procerus (Oberhauser) and Triadodiscus eomesozoicus (Oberhauser). A similar fauna was described by Benjamini (1988) in Israel, in levels dated as upper Anisian or lower Fassanian, possibly indicating a Sephardic origin and an older age for this group. Similar assemblages have been cited in the westernmost portion of the IR (Pérez-Arlucea and Trifonova, 1993), in the Pyrenees (Calvet

and Marzo, 1994) and in the Betic Ranges (Pérez-López et al., 2005). Interestingly, they are commonly associated with transgressive events. In the Calanda section, a highly diverse association was reported by Márquez-Aliaga et al. (1994) (Fig. 8). The presence of D. gerkey and S. oberhauseri, which are common in the Germanic area, and the Involutinidae (Aulatortus and Triadodiscus), which are also common in the Alpine domain, indicate clear influence of the Tethys realm in the Calanda area during the Ladinian. This association could be assigned to the T. mesotriasica Zone (A. praegaschei Subzone), indicating an upper Ladinan age. The assemblage suggests shallow, low energy environments with salinities higher than normal (Márquez, 2005). In the CCR, a foraminifer assemblage listed in Fig. 7, was described by Márquez et al. (1991) from the Rasquera Member of the Upper Muschelkalk. This assemblage is characterised by its relatively high diversity and clear influence of the Tethys realm with elements described from other basins of Europe and Asia (Márquez, 2005). Some of these foraminifers are shown in Fig. 17. 5.4. Conodonts The only Anisian assemblages found up to now in Iberia were reported and described by Márquez-Aliaga et al. (2000) from the Olesa and Vilella Baixa Members of the Lower Muschelkalk unit of the southern CCR (Fig. 6). These assemblages include Paragondolella bulgarica Budurov and Stefanov (Fig. 18: 2), P. hanbulogi Sudar and Budurv, P. bifurcate Budurov and Stefanov, Neogondolella constricta Mosher and Clark, (Fig. 18: 1), N. cornuta Budurov and Stefanov, N. excentrica Budurov and Stefanov, N. basisymmetrica Budurov and Stefanov, and Neogondolella longa Budurov and Stefanov. They indicate a middle– late Pelsonian to late Illyrian age and represent a typical Tethysian assemblage. Similar assemblages have been described in Bulgaria (Budurov et al., 1993), Italy (Mietto and Petroni, 1980) and Serbia (Urosevic and Sudar, 1991). The Ladinian record of conodonts encompasses the upper part of the Cañete Formation in the IR (both the Mediterranean Triassic domain and southern Levantine–Balearic Triassic domain), the Rasquera and Tivissa Members of the Upper Muschelkalk of the CCR (Figs. 6, 7, 8 and 10), and equivalent levels of the Calanda section (Plasencia, 2009). This record is characterised by a low species biodiversity: 90% of the samples are mono-specific restricted to Pseudofurnishius

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Fig. 13. Anisian cosmopolitan bivalves of Palaeotethys affinity. Landete Formation Iberian Range. 1. Myophoria vulgaris (Schlotheim). 2. Neoschizodus laevigatus (Goldfuss). 3. Neoschizodus orbicularis Bronn. 4. Neoschizodus ovatus (Goldfuss). 5. Elegantina sp. 6. Modiolus sp. 7. Burmesia posteroradiata Cox. 8. Pleuromya elongata (Schlotheim). 9. Unionites fassaensis (Wissmann). 10. Hoernesia socialis (Schlotheim).

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Fig. 14. Ladinian cosmopolitan bivalves, brachiopods and gastropods of Tethys affinity (Alpine domain), Cañete Formation Iberian Ranges. 1. Pseudocorbula gregaria (Munsteri). 2. Daonella cf. lommeli (Wissman). 3. Bakevellia costata (Schlotheim). 4, 5. Lingularia af. smirnovae (Biernat y Emig). 6. Neoschizodus laevigatus (Goldfuss). 7. “Natica” sp. 8. Modiolus myoconchaeformis (Philippi). 9. Umbostrea cristadiformis (Schlotheim). 10. Unionites munsteri (Wissman).

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Fig. 15. Ladinian bivalves of Sephardic affinity. Cañete Formation Iberian Range. 1. Pseudoplacunopsis teruelensis Wurm. 2. Limea vilasecai (Schmidt). 3. Gervillia joleaudi (Schmidt). 4. Costatoria aff. kiliani (Schmidt). 5. Gervillia joleaudi (Schmidt). 6. Elegantinia sublaevis (Schmidt). Scale = 0.5 cm.

murcianus Van den Boogaard or Sephardiella mungoensis (Diebel) (Fig. 18: 3–6), the former being more common. These two species were traditionally attributed to the Sephardic province. It should be noted, however, that recent studies have revealed a much wider

distribution than previously assumed. In fact, the original delimitation of the Sephardic Province (Hirsch, 1972) was primarily based on the distribution of P. murcianus in regions such as Israel (Huddle, 1970), Jordan (Bandel and Waksmundzki, 1985), Turkey (Nicora, 1981)

Fig. 16. Anisian (1–5) and Ladinian (6–10) foraminifers present in E Iberia. 1, 2. Hoyenella sinensis (Ho). 3. Turriglomina mesotriasica (Koehn–Zaninetti). 4. Paulbronnimannia judicarensis (Permoli–Silva). 5. Abriolina cf. mediterranea (Luperto). 6, 7. Lamelliconus ex gr. ventroplanus-biconvexus (Oberhauser). 8. Lamelliconus multispirus (Oberhauser). 9. Lamelliconus procerus (Liebus). 10. Nodosaria ordinata Trifonova.

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Tunisia (Rakus, 1981) and Sicily (Gullo and Kozur, 1991). However, later findings in the upper Ladinian and lower Carnian rocks in other areas of the Tethys, extended the realm of P. murcianus beyond the original Sephardic province to the Southern Alps of Italy (Nicora, 1981), Slovenia (Ramovs, 1977), Croatia (Balini et al., 2000), Serbia (Budurov and Pantic, 1973), and Hungary (Kozur, 1993), as well as southwestern China (Yang et al., 2001) and the Malayan Peninsula (Nogami, 1968; Ishida and Hirsch, 2011; Plasencia et al., in press). S. mungoensis was first described by Hirsch (1966) in the CCR near Mora de Ebro, although its geographical distribution was also very wide. Initially, during the early Longobardian, the species may have been limited to the Sephardic Province, i.e., Israel (Huddle, 1970), Palestine (Eicher and Mosher, 1974) and Jordan (Bandel and Waksmundzki, 1985). Later, during the middle and late Longobardian, its distribution extended to include areas as far away as Spain (Plasencia et al., 2007), Italy (Mastandrea et al., 1998), Bulgaria (Budurov, 1976), Russia (Klets, 1995), China (Buryi, 1996), and North America (Orchard and Balini, 2007). 5.5. Chondricthyans Detailed studies on Chondricthyans have only been carried out in the Ladinian (upper Cañete Formation) of the Mediterranean Triassic domain and southern part of the Levantine–Balearic Triassic domain of the IR (Figs. 6 and 10). The following species, shown in Fig. 18(7–13), have been described from these areas (Botella et al., 2009; Pla et al., 2013): Hybodus bugarensis (Pla, Márquez-Aliaga and Botella), Hybodus plicatilis (Agassiz), Lissodus aff. L. lepagei (Duffin), Palaeobates angustissimus (Agassiz), Prolatodon bucheri (Cuny, Rieppel, and Sander), Prolatodon contrarius (Johns, Barnes and Orchard) and Pseudodalatias henarejensis (Botella, Plasencia, Márquez-Aliaga, Cuny and Dorka). H. bugarensis and P. henarejensis have only been found in the IR, but other species, such as P. bucheri and P. contrarius, have also been described in North America and China (Chen et al., 2007). All these species are exclusively non-neoselachian sharks, confirming that by the Middle Triassic, neoselachians were rare in Europe. In terms of diversity, the chondrichthyan fauna were dominated by durophagous sharks, such as Lissodus aff. L. lepagei, Palaeobates angustissimus, Prolatodon bucheri and Prolatodon contrarius (Botella et al., 2009; Escudero-Mozo et al., 2012b; Pla et al., 2013). 5.6. Other fossils Gastropods abide at the top of the Cañete Formation in the IR (in the Mediterranean Triassic domain, specifically in the Calanda section and southern Levantine–Balearic Triassic domain). They are usually poorly preserved, and only “Natica” cf. stanenensis Pichl, Loxonema sp., Natica sp., Turbonilla dubia (Munsteri) and Zigopleura have been so far identified (Figs. 6, 8 and 10) (Márquez-Aliaga et al., 1994). Brachiopods are scarce in the Anisian and Ladinian of Iberia, with only one species described for the Anisian (Mentzelia mentzeli) in the southern CCR and in the Serra section of the northern Levantine–Balearic Triassic domain (Figs. 5B and 9) (Schmidt, 1932; López-Gómez et al., 1998). Coenothyris sp. and Lingularia cf. smirnovae (Biernat and Emig) specimens were described by Márquez-Aliaga et al. (1994, 1999) for the Ladinian units of the Mediterranean Triassic and southern Levantine–Balearic Triassic domains of the IR, as well as the Calanda section (Figs. 6, 8 and 10). 6. Palaeogeographic significance and discussion The recovery of marine fauna during the aftermath of the endPermian mass extinction was marked by pulses indicating alternating

favourable and unfavourable conditions determining an uneven process of recovery (Woods and Baud, 2008; Stanley, 2009). Such fluctuating conditions generated an immature, poorly functioning ecosystem that only with great difficulty could adapt to these extreme environmental changes (Tong et al., 2007). By the end of the Early Triassic or even beginning of the Middle Triassic, complete recovery had not begun in the western Tethys area and this gradual recovery process was indicated by increased tiering above and below the substrate (Twitchett, 1999; Márquez, 2005; Marshall and Jacobs, 2009). Although many of the palaeoenvironmental changes that took place at this time are most likely repeated worldwide, poor biostratigraphic resolution during this recovery period makes correlations difficult, even between contiguous basins. Rapid palaeogeographical changes in the Pangea supercontinent were an important factor controlling marine fauna distribution and this led to the different characteristics between basins during the Early Middle Triassic. Marine platforms suffered rapid changes during the different sea-level oscillations of the Middle Triassic. Microcontinents, such as Cimmeria in the Tethys realm, underwent permanent changes (Brühwiler et al., 2009). Some areas of the westernmost Tethys were exposed or covered by continental sediments until the Bithynian (Bourquin et al., 2011). Eastern Iberia was most likely first covered by the Tethys Sea during the Pelsonian (Galán-Abellán et al., 2013). During the Middle Triassic, Iberia experienced clear alternating continental-marine effects that gave rise to the particular evolution and distribution of its fauna. This detailed study of the main fossil assemblages of E. Iberia (IR and CCR) for this time interval with special emphasis on their affinities with other bioprovinces of the Tethys realm, reveals significant palaeobiogeographic differences between the Anisian and Ladinian associations. This information combined with the new stratigraphic data available, indicates that rapid palaeogeographical variations occurred during the Anisian–Ladinian transition in this area related to the evolution of the western Tethys. The Triassic palaeogeography of the western Tethys was clearly controlled by the intra-Pangea dextral shear, which separated the ancient Laurasia and Gondwana continents (Gutiérrez-Alonso et al., 2008; Muttoni et al., 2009). The shear alignment linked the southern borders of North America and Europe reaching the Palaeotethys between Iberia and the Morocco Massif (Fig. 19). Lateral movements of the shear structure gave rise to perpendicular transference fractures that controlled Cimmerian movements towards the north and therefore the development of the Palaeotethys and Neotethys (Ziegler and Stampfli, 2001; Stampfli and Borel, 2002). The general movement of Cimmeria in the Middle Triassic was related to Pangean rifting (Yin and Song, 2013). This movement was important in the westernmost Tethys, because this continent controlled the regional palaeogeography and therefore the different Middle Triassic marine incursions in this area, including Iberia, where two transgressive–regressive cycles took place in the middle–upper Anisian (Pelsonian–lower Illyrian) and upper Anisian–Ladinian. These marine incursions followed narrow corridors during the Anisian, controlling the development of different shallow marine environments and their fauna. The fauna examined in the different domains of Iberia draw a precise image of the distribution of these corridors and their dramatic changes from the middle Anisian to the early Ladinian. The data derived from the ammonoids, bivalves and foraminifers of the lowest T–R cycle indicate that Iberia was only connected with the Palaeotethys (Alpine/Germanic domains) during this time (Fig. 19). The absence of this cycle in the Levantine–Balearic Triassic domain indicates that this area constituted a topographic high, preventing possible marine incursions across the SE of Iberia during this time. Only the CCR, Calanda area and the central part of the IR (Mediterranean Triassic domain) where this cycle is recorded, formed a single palaeogeographic

Fig. 17. Selected Ladinian foraminifers present in the Rasquera Member of the Catalan Coastal Ranges. 1. Reophax asperus Cushman Waters. 2. Cyclogira pachygira (Gurmbel). 3, 4. Calcitornella sp. 5, 6, 7. Duostomina cf. alta Kristan–Tollman. 8, 9, 10. Oberhauserella mesotriasica (Oberhauser).

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marine corridor with a few small isolated highs while allowing for overflow of the Palaeotethys shallow marine waters with representative Alpine/Germanic bioprovince fauna for the first time. These fauna include Paraceratites and Olesites ammonoids specimens in the CCR, as well as cosmopolitan bivalves from shallow and restricted marine environments. During the late Anisian a new transgressive pulse (late Anisian– Ladinian age) is recorded in all the study areas, including the Levantine–Balearic Triassic domain. The Anisian fossil record of this cycle is scarce, but the bivalve assemblage found at the base of the Cañete Formation (Serra section) indicates, as in the previous cycle, a connection with the Palaeotethys and shallow restricted environments. However, the data derived from the facies and fossil assemblages indicate a significant change at the base of the Fassanian. This change is marked by the development of deeper environments (Rasquera Member and Henarejos Member), the presence of open marine bivalves (Daonella and Bositras), and by the appearance of ammonites Eoprotrachyceras curionii (Mojsisovics). Similar features have been described for other basins of the western Tethys such as the Betic Ranges (e.g. Pérez-Valera, 2005; Pérez-Valera and Pérez-López, 2008) or Minorca (Escudero-Mozo et al., 2014). Also, in association with this transgressive pulse, some ammonoid specimens, such as Gevanites archei (Goy) (Sephardic form), combined with other generalist forms of the Tethys realm reached Iberia during the late Fassanian. Most of the Ladinian genera of ammonoids in the study area (Eoprotrachyceras, Protrachyceras, Anolcites) broadly indicate their provenance from the Alpine realm, while the Gevanites genus (Hungaritidae) indicates a provenance from the Sephardic bioprovince (Hirsch, 1972, 1977; Parnes et al., 1985; Hirsch et al., 1987; Hirsch et al., 2014). Bivalve assemblages provide similar information to ammonites, since along with the cosmopolitan Tethyan bivalves of Alpine affinity several bivalves appear characteristics of the Sephardic bioprovince (Gervillia joleaudi, Pseudoplacunopsis teruelensis, Elegantina sublaevis, Limea vilasecai and Costatoria kiliani). No Shephardic ammonoids or bivalves reached the CCR during the Fassanian. These data demonstrate a mixed origin for the assemblages in the IR during the Fassanian, as supported by foraminifer, conodont, bivalve, and chondricthyan fauna. The conodont Pseudofurnishius murcianus, also represented across broad areas of the Tethys realm, is recorded in all Iberian domains including the CCR where no other fossils of the Sephardic bioprovince are present. In contrast, the ammonite, conodont and bivalve associations described from the Betic Ranges, situated to the southeast of E Iberia during that time, clearly belong to the Sephardic bioprovince (Hirsch, 1977; Hirsch et al., 1987; Goy, 1995; Márquez-Aliaga and Martínez, 1996; Pérez-Valera, 2005; Pérez-López and Pérez-Valera, 2007; Hirsch et al., 2014). The faunal distribution pattern indicates a clear increase in Sephardic faunal influence towards the south during the Fassanian. The general fossil content indicates a clear increase in diversity during the upper Fassanian–lower Longobardian. This increase reflects connection between sea corridors in the western Tethys realm since this period of time had a more humid climate (Sellwood and Valdes, 2007; Preto et al., 2010). This climate led to increased continental run-off and thus to the input of nutrients to marine waters, allowing for punctuated increases in faunal diversity (Algeo et al., 2011). This time period also witnessed the start of a general trend towards cooler waters, contributing to an increase in faunal diversity and more rapid recovery (Woods, 2005; Kiessling, 2010; Sun et al., 2012). Thus, during the Anisian–Ladinian transition, and mainly during the beginning of the Ladinian (Fassanian), marine connections in the

westernmost Tethys realm increased creating new and wider interconnected corridors (Fig. 19). On a regional scale, progressively larger areas of the deeply truncated Variscan orogen thermally subsided below the erosional base level and were transgressed by the eustatically rising Tethys (Ziegler and Stampfli, 2001; Vargas et al., 2009). The remnants of the Palaeotethys were almost closed when the Cimmerian microcontinent moved northwards and collided with the Variscan deformed Pelagonia block (Stampfli and Hochard, 2009). The combination of thermal subsidence, eustatic rises in sea-level and Cimmerian movements gave rise to a new palaeogeographic configuration in which the remnants of the Palaeotethys and the Neotethys eliminated geographical barriers and became widely connected (Fig. 19). Under this new palaeogeographic framework, the continental westernmost Tethys areas were rapidly invaded by marine waters in the late Anisian–early Ladinian, creating wide pathways approaching new western Pangea terrains. Iberia was affected by this transgressive episode and, for the first time, marine waters from different realms, in the absence of palaeogeographical barriers, invaded the eastern half of Iberia (Iberian Ranges, Catalan Coastal Ranges, and Betic Ranges) and the present-day Mediterranean Majorca, Minorca, Corsica and Sardinia islands, allowing for the migration of both the Sephardic and Alpine bioprovinces. 7. Conclusions This integrated study of the Middle Triassic carbonate platforms (Muschelkalk facies) of E Iberia (Iberian Ranges and Catalan Coastal Ranges) based on stratigraphic, sedimentologic and palaeontologic data unveils significant palaeobio- and palaeogeographical changes that serve to explain the evolution of the different palaeogeographic domains of Iberia. Based on these findings, a new palaeogeographic reconstruction of the westernmost Tethys is provided for this period. For the Middle Triassic three palaeogeographic domains of Iberia show deposition of Muschelkalk facies: the Iberian, Mediterranean and Levantine–Balearic Triassic. The Muschelkalk facies of theses domains record two broad transgressive–regressive cycles of the Tethys Sea, Pelsonian–early Illyrian and late Illyrian–Longobardian. The first incursion of the Tethys Sea is only recorded in the Mediterranean Triassic domain. This incursion reached the eastern part of Iberia during the Pelsonian through a small palaeogeographic marine corridor that crossed the southern CCR and only partially invaded the IR, while the rest of Iberia was a continentally elevated area. The fauna associated with this first incursion is typical of the Alpine/Germanic bioprovince related to the Palaeotethys. The upper Illyrian–Longobardian T–R cycle, represents a new eustatic sea-level rise that took place in a new palaeogeographic setting related to the intra-Pangea dextral shear and the northward movement of the Cimmerian microcontinent. This new eustatic sea-level rise caused a larger more generalised westward onlap of the Tethys sea onto vast continental lands, including the differentiated domains of Iberia, as well as Majorca, Minorca, Sardinia and Corsica, which were elevated lands during the late Anisian–Early Ladinian. The carbonate units of this new setting yield fossil assemblages (ammonites, bivalves, foraminifers and conodonts) of a mixed faunal origin and contain specimens from both the Alpine and Sephardic bioprovinces related to the Neotethys. The fauna of this latter bioprovince shows progressive northwards migration during the Ladinian, from the Betics to the Iberian Basin, but no Sephardic species are found in the CCR, except Longobardian conodonts. The fossil faunal record of Iberia indicates an increasing diversity during the upper Fassanian–lower Longobardian interval. This newly

Fig. 18. Middle Triassic conodonts (1–6) and chondrichthyans (7–13) in E Iberia. 1. Neogondolella constricta (Mosher and Clark). L'Ametlla, Tarragona, CCR, Anisian; 2. Paragondolella bulgarica Budurov and Stefanov, L'Ametlla, Tarragona, CCR, Anisian; 3. Pseudofurnishius murcianus van den Boogaard, Henarejos, IR, Ladinian; 4. Pseudofurnishius murcianus van den Boogaard, Henarejos, IR, Ladinian; 5. Pseudofurnishius murcianus van den Boogaard, Henarejos, IR, Ladinian; 6. Sephardiella mungoensis (Diebel), CCR, Ladinian; 7. Prolatodon bucheri (Cuny, Rieppel, and Sander). Henarejos, IR, Ladinian; 8. Paleobates angustissimus (Agassiz), Bugarra, IR, Ladinian; 9. Hybodus plicatilis Agassiz, Bugarra, IR, Ladinian; 10. Lissodus aff. L. lepagei. Bugarra, IR, Ladinian; 11. Pseudodalatias henarejensis Botella, Plasencia, Márquez-Aliaga, Cuny and Dorka, Henarejos, IR, Ladinian; 12. Prolatodon contrarius (Johns, Barnes, and Orchard) Bugarra, IR, Ladinian; 13. Hybodus bugarensis, Pla, Márquez-Aliaga and Botella, Bugarra, IR, Ladinian.

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257

Fig. 19. Palaeogeographic reconstructions of the westernmost Tethyan realm for the Middle Triassic: middle Anisian (Pelsonian) and the early Ladinian (Fassanian). The palaeogeographic map is modified from Ziegler and Stampfli (2001), Stampfli and Borel (2002) and Muttoni et al. (2009).

proposed palaeogeographic scenario suggests greater connections between sea corridors in the western Tethys realm, increased run-off and the consequent input of nutrients to marine waters. All this would

have induced episodes of punctual increases in faunal diversity and given rise to the onset of a period generally trending towards cooler marine waters.

258

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Acknowledgements This work was supported by a FPI predoctoral contract awarded to the first author and by projects CGL2008-00093 and CGL2011-24048, both funded by the Spanish Ministry of Economy and Competitiveness. The paper is also a contribution to the following research projects: “Sistemas Sedimentarios y Variabilidad Climática” (642853) of the CSIC, and Basin Analysis (910429), and Paleoclimatology and Global Change (910198) of the Universidad Complutense de Madrid. The authors thank Francis Hirsch and an anonymous reviewer, and the Editor Finn Surlyk, for their helpful comments on the manuscript.

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