Survival of early Viséan foraminifers in the Western Meseta of Morocco

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Survival of early Viséan foraminifers in the Western Meseta of Morocco Pedro Cózar a,∗ , Daniel Vachard b , Alain Izart c , Ismael Coronado d a

Instituto de Geociencias CSIC-UCM, Ciudad Universitaria, 28040 Madrid, Spain b 1 rue des Tilleuls, 59152 Gruson, France c 202 Chemin de Cabanis, 34730 Prades-le-Lez, France d Institute of Paleobiology, ul. Twarda 51/55, PL-00-818 Warsaw, Poland

Received 21 March 2019; received in revised form 14 May 2019; accepted 3 June 2019

Abstract Typical early Viséan foraminiferal assemblages documented in outcrops of the Western Meseta of Morocco led to the erroneous biostratigraphic dating of areas that later, after much effort, have been demonstrated to correspond to the mid and late Viséan. These sections are analyzed to decipher if they are formed by reworked specimens or if this fauna really survived into younger rocks. Key sections are located in vast areas of the Western Meseta, in the north of the Azrou-Khenifra basin, Fourhal area, El Hammam Ridge, Oulmès area, Sidi Bettache basin (all on the north of the Western Meseta), and the Skoura region (on the southern border). The environmental settings as well as the taphonomical stage of preservation of the foraminifers suggest that most of the assemblages are composed of autochthonous or parautochthonous foraminifers and, thus, that the fauna really survived into the middle and even upper Viséan carbonate platforms of the Western Meseta. In some sections, the early Viséan foraminifers share the same levels with mid Viséan foraminifers; in others, they share the same levels with mid and late Viséan foraminifers, and in a third group, the early Viséan foraminifers occur in the same stratigraphic sections with mid and late Viséan assemblages, although never in exactly the same stratigraphic levels but alternating. These distributions exemplify the three patterns of interaction between the stratigraphic and biogeographic ranges of the assemblages. The absence of similar patterns is noteworthy in the surrounding regions of the Palaeotethys, and thus, this anomalous distribution can be discarded as a matter of dispersal from neighbouring regions. The absence of similar patterns in basins situated far from the Western Meseta allows abiotic factors to be discarded, such as tectonic/environmental setting, palaeolatitude or isolation. The most plausible hypothesis to explain the survival of these fauna is related to biotic factors, such as species interaction and competition, and they can be compared to some similar modern ecological patterns. However, the primary triggering factor allowing these biotic factors to interact is considered to be the late arrival of the mid Viséan foraminifers to the Western Meseta, allowing them to occupy niches completely different from the rest of the Palaeotethys. © 2019 Elsevier Ireland Ltd Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. Published by Elsevier B.V. All rights reserved. Keywords: Foraminifers; Mississippian; Species-interaction; Long-time persistence; Northern Africa

1. Introduction The early Viséan in Morocco has been a matter of discussion from the early studies in the country (e.g., Termier, 1936), where its occurrence has been questioned in vast areas of the Western Meseta of Morocco (Tahiri, 1991). Due to the predominance of shallow-water platforms, the study of foraminifers was

∗

Corresponding author. E-mail addresses: [email protected] (P. Cózar), [email protected] (D. Vachard), [email protected] (A. Izart), [email protected] (I. Coronado).

vital in the improvement of the regional biostratigraphy in most basins, which resulted in a regional zonal scheme for Morocco (Vachard, 1988; Vachard and Tahiri, 1991). The zonations contains up to 13 zones, from Cfm1, in the lower/middle Viséan boundary interval, to Cfm9 for the Viséan (Fig. 1), and the Cfm10 to Cfm13 for the Serpukhovian and the early Bashkirian. This biozonation reveals a late arrival of some Western European markers to Morocco, mostly in the lower zones, where the main markers for the mid Viséan in Europe, Pojarkovella nibelis and Koskinotextularia bradyi, first occur near the top of the mid Viséan in Morocco. Meanwhile, the European mid Viséan Pojarkovella nibelis is only known in one Morocco section for

https://doi.org/10.1016/j.palwor.2019.06.001 1871-174X/© 2019 Elsevier Ireland Ltd Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. Published by Elsevier B.V. All rights reserved.

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Fig. 1. Stratigraphic correlation between the foraminiferal zonal schemes in Belgium and northern France (based on Conil et al., 1991; Poty et al., 2006); the British Isles includes zones defined by Conil et al. (1980); and the Moroccan biozones and markers (Vachard and Tahiri, 1991; Izart et al., 2017).

this substage (Vieslet, 1983), whereas the rest of specimens documented from Morocco occur in the late Viséan (Izart et al., 2001). To complicate the zonation further, the basal zone of this zonal scheme, the Cfm1, was never described clearly as early or mid Viséan due to the mixture of foraminifers, spanning assemblages in the lower/middle Viséan boundary interval in Western Europe. Furthermore, European chronozones defined for Belgium and France were the V1, equated to the early Viséan, V2 as mid Viséan and V3 as late Viséan (Conil and Lys, 1964). Later, Conil et al. (1977) considered the foraminiferal subzone equivalent to the V2a as part of the early Viséan, and the equivalent of the V3a part of the mid Viséan (Fig. 1). Stratigraphic sections dated as early Viséan (or V2a subzone) are common in Morocco based on the occurrence of Uralodiscus, Glomodiscus, Propermodiscus or Rectodiscus assemblages (e.g., Verset, 1988; Izart et al., 1989; Charrière, 1990; Huvelin and Mamet, 1997). However, later studies in the same sections have shown also the occurrence of mid or late Viséan markers (e.g., Berkhli, 1999). These assemblages have been never studied from a taphonomical point of view to elucidate if they are reworked or if they survived for a longer time in Morocco. Nevertheless, it must be noted that these unusual early Viséan assemblages are restricted to basins in the Western Meseta of Morocco (Fig. 2), because there is no lower and middle Viséan carbonates in the Eastern Moroccan Meseta (Berkhli, 1993), and they are not recorded in the upper Viséan limestones at Jerada (see Vachard and Berkhli, 1992; Berkhli et al., 1999). Similarly, in the Sahara basins (Cózar et al., 2016), there is no evidence of early Viséan foraminifers in the mid or late Viséan (Sebbar et al., 2000; Cózar et al., 2014). The main aim of this study is to demonstrate that foraminifers, typically distributed in the early Viséan in Europe, survived during the mid and late Viséan of the Western Meseta of Morocco. This implies the low reliability of some dating published in Morocco, and, thus, more detailed studies are necessary at some outcrops. Furthermore, the possible palaeobiogeographic implications or causes for these unusual stratigraphic records are discussed.

Fig. 2. Main Carboniferous outcrops in Morocco. Abbreviations: MMFZ – Middle Meseta Fault Zone, RTFZ – Rabat-Tiflet Fault Zone, SAF – South Atlas Front, SMF – South Meseta Fault, WMSF – Western Meseta Shear Fault.

2. Geological setting of the Moroccan Meseta The Moroccan Meseta is composed of several structural blocks that correspond to perigondwanan terranes of restricted displacements in relationship with northwestern border of Gondwana. Some of these blocks, including Sehoul Block, Eastern Meseta and Western Meseta, juxtaposed during the late Devonian–Pennsylvanian (Hoepffner et al., 2005; Michard et al., 2010). Each block has a different tectonics, metamorphism and stratigraphic succession, mostly for that of the Sehoul Block, which occurs north of the Rabat-Tiflet Fault Zone (RTFZ) and extends down of the Rif foredeep (Fig. 2). This difference is attributed to a distinct origin of the terrane, because it could be derived from Avalonia (Simancas et al., 2005) or from Meguma (Michard et al., 2010). The Western Meseta is also subdivided in regions: the Central Zone, Nappes Zone, Coastal Block, the western Meseta Shear Zone (a thin band limited between the Coastal and Central blocks), and the sub-Meseta or High Atlas

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Zone in the south (limited between the Southern Meseta Fault and the South Atlas Front) (Fig. 2). Three tectonostratigraphic events are usually considered for the Variscan orogen as a response to three tectonic phases (Michard et al., 2010): An Eovariscan deformation (372–366 Ma) well represented in the Western Meseta, particularly in the Nappes Zone, Coastal Block and also in the Sehoul Block. A second intra-Viséan deformation (330–320 Ma), which occurs in the Nappes Zone and the Eastern Meseta. A final Pennsylvanian deformation (300–290 Ma), well represented in the Western and Eastern Meseta. Hoepffner et al. (2005) highlighted that isotopic ages suggest a continuous and progressive regional shortening, more developed in the northern basins than in the southern outcrops. The analysed outcrops are located in the Central Zone of the Western Meseta, where Carboniferous basins are rather common (Fig. 2). There is a predominance of turbidites filling the centre of each basin with a variable shelf deposits in the borders, siliciclastics as well as carbonates. Several geodynamic models have been proposed for the formation of the Carboniferous Moroccan Meseta: (i) an oceanic subduction followed by continental collision, with the subduction plane dipping to the west or to the east (below the Meseta and Sahara respectively); (ii) a limited continental subduction along crustal faults dipping to the east or to the west (see Izart et al., 2017 and references therein); (iii) a combined system of tensional forces and mantle plume convection leading to a short distance dispersal of the terranes, generating an archipelago with a network of seaways (Crowley et al., 2000); and (iv) a dextral strike-slip in the South Atlas Front with a Pennsylvanian compressional collision (Simancas et al., 2005). Therefore, most authors agree that the Moroccan Meseta is the result of an intracontinental evolution during the Carboniferous. The Sidi Bettache, Rehamna and Jebilet basins are interpreted as pullapart basins during a transtensive context (Piqué, 1979; Essaïfi, 1995; El Kamel, 2004), whereas the Adarouch Khenifra basin started in a similar way but it evolved in a transpressive foreland (Bouabdelli and Piqué, 1996). 3. Early Viséan foraminifers in the Western Meseta The typical fauna of the early Viséan is recorded in some regions of the Western Meseta, from northeast to northwest: Azrou-Khenifra, Fourhal, Oulmès-El Hamman Ridge and Sidi Bettache. In the south, only the Skoura boutonnière of the High Atlas (Fig. 2) records the lower Viséan, whereas in other outcrops of the High Atlas like Jebilets and Rehamna (Fig. 2), upper Viséan limestones occur. The mixture of the fauna from these areas has not been published (Izart, 1990; Vachard et al., 1991). 3.1. Azrou-Khenifra The most classic outcrop of the early Viséan of Morocco is El Goulib hill (Termier et al., 1975), but it is also the origin of the paradigm of the out-of-zone foraminifers in the Moroccan Meseta. Since the early publications, the succession was assigned to the V2a at the base and the V2b/V3 in the middle

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and upper parts (Termier et al., 1975; Vachard et al., 1977). The V2a assemblage contains numerous taxa that occur throughout the Viséan, which does not give us any substantial information. However, it also contains Lapparentidiscus nanus, Septabrunsiina minuta elegantula and Uralodiscus elongatus, which were genera extinct in Western Europe at the top of the early Viséan (Conil et al., 1980, 1991; Poty et al., 2006). Other assemblages recorded by the authors, and assigned to the V2b/V3, are the usual assemblages for the mid Viséan and even late Viséan. The occurrence of the most evolved representatives of Glomodiscus is not unusual, as well as Uralodiscus, characterized by the reduction of the contraforts and microgranular layer, because they are occasionally recorded up to the late Viséan (Conil, 1963, Conil and Lys, 1964; Vachard, 1977; Cózar, 2000; Brenckle, 2004; Somerville and Cózar, 2005; Vachard et al., 2016; Zandkarimi et al., 2016, 2017), and even these taxa are commonly recorded at the base of the Russian Tulian substage (equivalent to the middle Viséan) (e.g., Reitlinger et al., 1996; Brenckle and Milkina, 2003). Furthermore, it must be noticed that most of the typical mid Viséan fauna has extended stratigraphic ranges into the late Viséan (e.g., Conil et al., 1991), and the latter is recognized by the addition of new foraminifers. Important foraminiferal markers in the V2b assemblages in El Goulib also include Endothyranopsis compressa (a marker for the mid Viséan for Lipina and Reitlinger, 1970; Conil, 1977; Vachard, 1977; Kalvoda, 2002; Kulagina et al., 2003), and large Archaediscus chernoussovensis (at concavus stage). This evolutionary stage is considered to be a marker for the mid Viséan (Conil et al., 1980, 1991), so the entire assemblage should be considered at least as mid Viséan from the base. It must be noted that the authors did not used the term “early Viséan” for those V2a assemblages and, furthermore, they indirectly dealt with the outcrops as mid Viséan, as well as most of the outcrops between Oulmès and Agouraï, following the criteria of Termier (1936), who did not consider the existence of the early Viséan in this region following the traditional chronozones of Belgium and northern France (before the revision by Conil et al., 1977). Berkhli (1999) sampled up to 36 levels at El Goulib, of which the foraminiferal assemblages of 4 levels were published, but considered a reversed section. From the basal horizon, Archaediscus at concavus stage occurs, confirming that the section should be considered at least mid Viséan from its base. In samples Gb24 and Gb27, Uralodiscus ex gr. rotundus and Glomodiscus oblongus occur. In this case, some specimens are reworked (e.g., Berkhli, 1999, pl. 1, fig. 7), showing incongruous internal infilling and truncational facets (cf. Cózar, 2003a, 2005). However, the time interval of this reworking cannot be estimated because it could be from nearby beds where the taxon was well preserved, and, thus, geologically negligible. In the upper part of the succession, Archaediscus ex gr. karreri was recorded and thus, this part of the section was assigned to the Cf6␣. There is an alternation of limestone, sandstone and shale in the succession. The microfacies are packstone-grainstone with bioclasts (foraminifers, crinoids and bryozoans), ooids and round quartz. Hummocky cross-stratification and limestonesandstone couplets (Kelling and Mullin, 1975) are interpreted

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as storm and turbidite deposits above and below storm wave base on the outer platform (according to Walker, 1984, fig. 21). These main processes of storms and turbidites contribute to the degree of allochthony of the assemblages, forming taphofacies where autochthonous assemblages s.s. are absent, with important (10–100%) resedimented/transported specimens, but also usually with low percentages of reworked specimens (10–40%) (Cózar, 2003b), which is in agreement with the analyzed material. Consequently, foraminiferal assemblages from El Goulib are composed of typical mid Viséan foraminifers with sparse, supposedly “early Viséan” foraminifers, mostly transported from the platform facies where they were living. Ouarhache (1987) recorded the conodont Gnathodus girtyi girtyi, a taxon which extends rarely from the lower part of the early Asbian (Cf6␣–␤) up to the Namurian (Higgins, 1985; Varker and Sevastopulo, 1985). This coincides with the occurrence of some foraminifers assigned to the Cf6␣ at the top of the succession by Berkhli (1999). This suggests that: (1) the section should be mostly dated as mid Viséan to late Viséan in its upper part (Fig. 1), and (2) the “early Viséan” foraminifers survived in the unpreserved platform up to the late Viséan, and they concentrated in the turbidites of the same period (see Cózar, 2005). Vachard et al. (1977) recorded some beds with typical V2a foraminifers. In contrast, in the assemblages assigned to the V2b or V2/V3, they considered that some early Viséan foraminifers remained in the assemblages. Some of the most primitive taxa recorded in the latter assemblages are Forschia subangulata, Omphalotis exilis, Endothyra ex gr. similis, and Planoarchaediscus spirillinoides, which are well-known currently to extend through the Viséan, and even into the Serpukhovian. One of the most anomalous occurrences is that of Glomodiscus sp., but, as was previously discussed, rare specimens of this genus readily survived much later than the early Viséan. Stratigraphic levels described by Berkhli (1999) and Berkhli et al. (2000) contain some assemblages that were assigned to the V3b, whereas other stratigraphic levels contain assemblages assigned to the V2a/V2b. The foraminiferal assemblages in El Goulib thus seem to be excluding each other, and they overlapped in time, but their palaeogeographic ranges did not overlap, and remained separated in the same region. It must be also highlighted that the section was measured in an olistolite, because the surrounding shales contain the bivalve Posidonia becheri (Cf6␦ = Brigantian; Fig. 1) (Ouarhache, 1987). Reworked assemblages were identified in boulders of the conglomerates of the M’Taoutoult Formation (Verset, 1983; Ouarhache, 1987; Ouarhache et al., 1991), which was considered Namurian (using macroflora; Broutin in Ouarhache, 1987). Similar results in the Azrou-Khenifra basin are recognized in the Imouzzer du Kandar boutonnière. Charrière (1989, 1990) recorded an assemblage composed of Glomodiscus sp., Ammarchaediscus sp. (= Conilidiscus p.p.), Rectodiscus sp. (= Uralodiscus p.p.) and Archaediscus at involutus stage. All the specimens came from a single stratigraphic level. He assigned the entire succession to the V2a (as early Viséan). Ouarhache (1987) sampled nine horizons in the same lime-

stone ridge, among which, five levels contain typical mid Viséan foraminifers, and Glomodiscus miloni (as G. rigens) occurs in each sample. The specimens show the final whorl broken, with congruous cements and infillings with the matrix, which suggests that they could be transported, but not reworked. Other foraminifers in the assemblages allowed the dating of this locality as V2b–V3b (mid to late Viséan). Later, Berkhli (1999) considered the succession as V3b␥ due to the occurrence of Endothyranopsis ex gr. crassa and Koskinobigenerina prisca, which are markers of this age (Conil et al., 1980; Somerville, 2008). However, there are no typical “early Viséan” foraminiferal survivors in the assemblage listed by this author, apart from the mentioned Glomodiscus miloni. This absence is interpreted as depending on the sampled horizon. It is possible to find a typical late Viséan foraminiferal assemblage locally, as some sparse “early Viséan” foraminifers might have also survived, but generally in separated stratigraphic levels. 3.2. NW Fourhal This region is mostly composed of flysch facies of the late Viséan to early Bashkirian (Ghfir, 1993), although in the northwestern parts, between Agouraï town and the El Hamman Ridge, there are some sandy limestones that were attributed to the early to late Viséan (Berkhli, 1999; Berkhli et al., 2000). At the base of the succession, conglomerates and sandstones were considered as early Viséan due to some lateral outcrops of similar age. These outcrops contain a typical “early Viséan” foraminiferal assemblage: Glomodiscus oblongus, Uralodiscus sp. and Eotextularia diversa. These forms occur in the longest stratigraphic section, Butte 826, also known as Bou Alzaz. Higher levels in the succession contain horizons with typical “early Viséan” foraminifers such as Eoparastaffella, Glomodiscus oblongus, Conilidiscus and Paraarchaediscus (at involutus stage). In contrast, other stratigraphic horizons contain typical late Viséan foraminifers with sparse Glomodiscus and Archaediscus at involutus stage, and, finally, other levels only contain typical late Viséan foraminifers. In this stratigraphic section, the record of Conilidiscus, Uralodiscus and Glomodiscus up to the late Viséan is usual. The stage of preservation of the illustrated material is good: only some final whorls are broken, but infillings and matrix are congruous. Truncational facets are also observed (i.e., Conilidiscus cf. bucculentus in Berkhli et al., 2000, pl. 1, fig. 15), although the percentage of reworking cannot be estimated in the scarcely illustrated material. This Bou Alzaz section has been resampled from the basal conglomerates and sandstones (with local limestone lenses). Samples from the lower siliciclastic interval contain rare Archaediscus at concavus stage, so according to Conil et al. (1980, 1991) the succession should be considered as mid Viséan from its base, and late Viséan in the middle and higher parts. Specimens of varied species of Uralodiscus and Glomodiscus, as well as Eoparastaffella, in particular Eoparastaffella macdermoti, as well as Endolaxina and Pseudoplanoendothyra sp., are commonly recorded up to the late Viséan (Fig. 3). These assemblages occur predominantly as mixed assemblages, and are assigned to the late Viséan due to the occurrence of Archaediscus

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Fig. 3. Selected taphonomical processes in the Bou Alzaz section, scale bar = 200 ␮m. (A) Glomodiscus sp., remobilized and preserved inside an Earlandia vulgaris, level Pc 3217 (late Viséan). (B, C) Continuous distortion due to lithostatic pressure in Spinolaxina sp., levels Pc 3207 and 3209 (mid Viséan). (D) Crushed Uralodiscus or Conilidiscus, level Pc 3208 (mid Viséan). (E) Crushed Uralodiscus or Conilidiscus, level Pc 3210 (mid Viséan). (F) Glomodiscus sp. (cf. G. miloni) with periphery destroyed by pressure-dissolution (see Cózar, 2003b), level Pc 3215 (late Viséan). (G) Uralodiscus inflatus, with slightly deformed final whorl, level Pc 3215 (late Viséan). (H) Glomodiscus rigens, with the final whorls broken, level Pc 3216 (late Viséan). (I) Uralodiscus sp., with the final whorl broken, level Pc 3217 (late Viséan). (J) Reworked Archaediscus at angulatus stage, with a truncational facet that nearly eroded half of the specimen, level Pc 3217 (late Viséan). (K) Reworked Archaediscus at concavus stage, with a truncational facet and bioerosion in the periphery of the test, level Pc 3207 (mid Viséan). (L) Reworked Uralodiscus rotundus, with truncational facets and incongruous infilling, level Pc 3224 (late Viséan).

at angulatus stage, Pseudoendothyra struvei and Endothyranopsis crassa, and also Glomodiscus, Uralodiscus and Archaediscus at involutus stage. This simultaneous distribution contrasts to outcrops in the Azrou-Khenifra basin, and in Agouraï, where assemblages are not excluding each other, and survivals of the early Viséan were co-occurring with the late Viséan fauna, and thus, sharing stratigraphic and palaeogeographic ranges. The succession in Bou Alzaz is composed of an alternation of limestone, sandstone and shale. Sandstone with crossstratification of low angle was deposited on a beach during the Devonian. The limestones and sandstones present couplets and cross-stratifications interpreted as storm deposits on a platform (Berkhli et al., 2000). Many of the specimens are well preserved, with intact tests or diagenetic continuous distortions, but final broken chambers

(typically in endothyrids) as well as final broken whorls (in archaediscids) occur. Both facts are interpreted as a result of transport or remobilization, but not reworking. Crushed tests are also recorded, which is interpreted as caused by lithostatic pressure on non-cemented specimens, and thus, is also indicative of parautochthonous assemblages (Cózar, 2005) due to the general remobilization of the storm deposits (Cózar, 2003c). Exceptionally, truncational facets are recognized, which demonstrate the occurrence of sparse reworked specimens, although the percentage of these specimens is negligible (Fig. 3). 3.3. Oulmès and El Hammam Ridge The outcrops northeast of Oulmès town are related to the Bouqachmir Formation located north of the Oulmès town

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and Jbel Moufras Formation located on the El Hammam Ridge, which extend up to the late Viséan. There, species of Archaediscus at involutus stage, Glomodiscus and Eotextularia are common (Tahiri, 1991). Furthermore, Lapparentidiscus bokanensis, Pseudolituotubella, Glomodiscus lenitortus and Pseudoplanoendothyra also occur in the late Viséan (Vachard and Tahiri, 1991). None of the illustrated material shows any evidence of reworking. Apparently, there is a mixture of typical early and mid Viséan foraminifers, whereas in higher levels of the succession (late Viséan), only Glomodiscus cf. oblongus persisted, but rarely. This is interpreted as the mixture of fauna that can be only possible in the lower part of the succession, whereas the fauna recorded in higher levels is the typical taxa that survived elsewhere in the Palaeotethys. Izart et al. (2001) published assemblages from Jbel Moufras Formation, including Glomodiscus lenitortus, Lapparentidiscus bokanensis, Pseudoplanoendothyra, Uralodiscus rotundus, Uralodiscus elongatus and Uralodiscus sp. in levels of the mid Viséan. These assemblages do not show traces of reworking. Similarly to the Bouqachmir Formation, there is a co-existence of fauna only during the mid Viséan, but not in the late Viséan. The Bouqachmir Formation consists of an alternation of turbidites, calciturbidites, and olistolites of platform limestone and shale deposited in a marine basin, which was a prolongation of the Sidi Bettache basin towards the east. A grainstone bed with ripples is known in the Sidi Abbou syncline with ooids and bioclasts (foraminifers, crinoids, brachiopods and algae). It is interpreted as a storm deposit on a platform located at the northern border of the Oulmès Ridge. The Jbel Moufras Formation contains limestone and shale deposited on a marine platform on the El Hammam Ridge. The limestones are wackestone and packstone with foraminifers, algae, crinoids and bryozoans. They were deposited in the inner and middle platform on the El Hammam Ridge (Izart et al., 2001). In all the studied sections in this region, it seems to be usual to have these faunas up to the mid Viséan, although they do not seem to extend into the late Viséan. 3.4. Sidi Bettache In the northeast of the basin, the best example of mixed faunas is well known in the south of the town of Tiflet (Vieslet, 1983). This outcrop is limited to a 2 m thick limestone and it was assigned to the mid Viséan (Cf5) because it contains all the typical European markers, Pojarkovella nibelis, Koskinotextularia bradyi, Endothyranopsis compressa (misidentified as E. crassa) and Archaediscus at concavus stage. The limestone lens rests on sandstone and conglomerate with siliceous pebbles. The limestone is a bioclastic packstone to grainstone rich in foraminifers, algae and corals deposited on the middle platform north of the Sidi Bettache basin. The foraminiferal assemblage also contains: Brenckleites fragilis (as Mediocris sp.), Conilidiscus sp. (as Rectodiscus), Conilidiscus settlensis, Cribranopsis foeda, Endolaxina laxa, Eotextularia diversa, Glomodiscus sp., Glomodiscus miloni, Glomodiscus oblongus, Glomodiscus praeconvexus, Lapparentidiscus sp., Latiendothyranopis aff. menneri solida,

Pseudolituotubella sp., Pseudolituotubella tenuissima, Spinobrunsiina sp. and Uralodiscus rotundus. All those species are widely known to have disappeared at the top of the lower Viséan in Western Europe, and their survival into the base of the mid Viséan in Eastern Europe is rare. The assemblages are well preserved, with rare broken final chambers or whorls and crushing, suggesting that they are mostly autochthonous assemblages, and, thus, early and mid Viséan foraminifers inhabited the same platform, sharing also their palaeogeographic ranges. In western outcrops of the Sidi Bettache area, this mixture of fauna is not observed, although middle Viséan limestones are rather scarce except in Sidi Radi and Sidi el Kacem with limestone-sandstone couplets interpreted as storm deposits in the outer and middle platform (Izart and Vieslet, 1988; Izart, 1990). The Sidi Sebaa section, in the Mdakra Massif, south of Sidi Bettache, is the only locality where the early Viséan fauna is recorded as restricted to lower Viséan limestones (Vachard and Fadli, 1991). There, only Glomodiscus oblongus extends above the first occurrences of Archaediscus at concavus stage, whereas other species of Glomodiscus, Uralodiscus or Pseudoplanoendothyra, disappeared at this level. There is a predominance of shales, with rare limestone beds, commonly in beds 5–30 cm think. Limestones are poorly bioclastic mudstones, bioclastic wackestones, with abundant brachiopods, bivalves and bryozoans, and sandy bioclastic wackestones, and, in the upper part of the succession, oolitic grainstones with cross-lamination are recorded. Environments are attributed to the middle to inner platform. 3.5. High Atlas In the southern border of the Meseta, in the High Atlas, the Skoura boutonnière contains a succession with early Viséan foraminifers (Izart et al., 1989). There, Glomodiscus oblongus, Glomodiscus miloni, Glomodiscus praeconvexus, Uralodiscus rotundus and Eotextularia diversa occur. Furthermore, Conilidiscus settlensis, Eoparastaffella vdovenkoae, Lapparentidiscus nanus, Latiendothyranopsis menneri solida, Conilidiscus bucculentus and Uralodiscus elongatus are also recorded. However, Archaediscus at concavus stage has been found in the oldest limestone beds in a re-sampling of the Assif N’Tanzouzmine section, the longest lower Viséan stratigraphic section in the region (Izart et al., 1989). The latter suggests that the succession is younger than was supposed, and it should be assigned to the mid Viséan. There is an alternation of sandstones and sandy limestones rich in crinoids (mostly in the lower part of the succession) with brachiopods, bryozoans, ostracods and foraminifers, and locally gastropods. Solitary rugose corals are abundant in the shales, which is the predominant lithology in the lower and upper part of the succession. They are autochthonous and paraautochthonous assemblages deposited on inner and middle platforms. The state of preservation is relatively better than in previous outcrops, and rarely, broken final whorls occur, suggesting generalized autochthonous assemblages. However, one level (Pc 5468) contains many reworked specimens, and nearly simi-

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Fig. 4. Selected taphonomical processes in the Assif N’Tanzouzmine section, scale bar = 200 ␮m. (A) Uralodiscus rotundus, with the final whorl broken, level Pc 5468 (mid Viséan). (B) Crushed Conilidiscus settlensis, with multiple perforations by bioerosion, level Pc 5468 (mid Viséan). (C) Broken final whorl in Uralodiscus rotundus with congruous cement, level Pc 5468 (mid Viséan). (D) Uralodiscus elongatus, with the final whorl broken and a congruous cement as the continuation of the external cement, level Pc 5483 (mid Viséan). (E) Reworked Uralodiscus elongatus, with truncational facet, level Pc 5468 (mid Viséan). (F) Reworked Uralodiscus sp., with truncational facet, eroded final whorl and incongruous infilling, level Pc 5468 (mid Viséan). (G) Uralodiscus elongatus, with the final whorl broken with apparent congruous cement and incipient development of a coating in the periphery; the specimen is interpreted as reworked because the development of the external cementation is before the break of the test, level Pc 5468 (mid Viséan). (H) Reworked Uralodiscus rotundus, with cement crust in the periphery and truncational facet, level Pc 5468 (mid Viséan). (I) Deformed Uralodiscus sp., with bioerosion; note that the micritic infilling inside the final whorls is intact in many parts of the final whorl, which is partly eroded by several truncational facets, level Pc 5468 (mid Viséan).

lar percentages of specimens without evidence of reworking (Fig. 4). The occurrence of this level does not minimize the fact that the overall assemblages in this locality should be considered as autochthonous, and some tempestite levels as para-autochthonous. This locality can be compared with Tiflet, where both faunas (that of the early and mid Viséan) shared the same ecological niches on the platform, in time and space. The assemblages of the Skoura boutonnière can be assigned to the basal assemblages of the mid Viséan, and, in contrast, the assemblages of Tiflet might be correlated with younger levels in the mid Viséan because all the foraminiferal guides are present. 4. Possible surrounding regions of influence In the Sahara, south of the Southern Atlas Front (Fig. 2), lower Viséan limestones are recorded on the southern flank of the Tindouf basin (e.g., Sebbar, 2000; Sebbar et al., 2000) or Tafilalt (Aretz et al., 2013; Mottequin et al., 2017), although no foraminifers have been described. Nevertheless, in the mid Viséan of the Tindouf basin, there is no survival of those forms (Sebbar et al., 2000), so this effect of longest persistence does not seem to occur in the Saharan basins. East of the Middle Meseta Fault Zone, in the eastern Meseta (Fig. 2), there are no lower and

middle Viséan limestones, and those supposed “early Viséan” foraminifers are not described in the late Viséan (Vachard and Berkhli, 1992; Berkhli et al., 1999). In Newfoundland and Nova Scotia in Canada, similarly, there are only representatives of the late Viséan foraminifers, and no “early Viséan” taxa were described (von Bitter et al., 2007). In SW Spain, in the Ossa-Morena Zone, mostly upper Viséan limestones are known (e.g., Cózar, 2003c), and apart from rarely evolved Glomodiscus rigens and Uralodiscus adindanii, no other primitive forms occur. Early Viséan foraminifers have been recently discovered in western outcrops of this zone, in the Cabrela basin in Portugal, typically composed of Uralodiscus, Glomodiscus and Archaediscus at involutus stage (J. Kalvoda, personal communication), but there, no mixture of fauna is recognized. A similar fauna occurs in the El Pimpollar outcrop, in the South Portuguese Zone, also in SW Spain. Boulders of limestones preserved in the andesitic-dacitic member contain assemblages with Glomodiscus miloni and Uralodiscus rotundus (Díez Montes et al., 2006). Other boulders may contain mid Viséan assemblages, but the foraminiferal faunas do not seem to be mixed. Simancas (1983) considered the calcareous beds as late Viséan, although he did not indicate precisely the

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levels where the fossils were found. Due to the fragmentary nature of the outcrop, it cannot be elucidated if the early Viséan foraminifers survived into the late Viséan, or, likely, it is an effect of the mixture of limestone blocks after the destruction of the unpreserved platform. The absence of a longer persistence of the early Viséan foraminifers in surrounding regions with a possible direct palaeobiogeographical communication with the Western Meseta of Morocco, allows rejection of this possible influence, and that those surviving foraminifers can be considered as restricted to the Western Meseta, and predominantly, in its northern part. Within this northern part, the regions situated in eastern positions show the persistence of the fauna up to the late Viséan, whereas the northwestern areas only extended up to the mid Viséan. In contrast, in the southern outcrops, the Skoura boutonnière might only represent the basal part of the mid Viséan due to the occurrence of Archaediscus at concavus stage, because the other more typical mid Viséan markers (Pojarkovella nibelis, Koskinotextularia bradyi, Endothyranopsis compressa) are not recorded, and the mid–late Viséan assemblages in the region (Jenny, 1988; Izart et al., 1989; Vachard et al., 1991) do not contain these primitive forms. As a consequence, they survived for a short time interval, rather different to the north of the Western Meseta.

5. Worldwide comparison Some of the more evolved species of the genera Uralodiscus and Glomodiscus are relatively common during the mid or late Viséan. However, more primitive forms are rarely documented in these substages. Apart from the afore-mentioned taxa, early Viséan foraminiferal survivals are only documented in the mid Viséan around the World. However, they have not been documented in the late Viséan anywhere other than in the sections of the northeastern Western Meseta. These unusual taxa in the mid Viséan around the World have been apparently documented in Ireland (Austin et al., 1973), Belgium (Conil, 1963; Kimpe et al., 1978), Moravia (Kalvoda, 1983, 1990, 2002), France (Vachard et al., 2018) and China (Devuyst et al., 2003; Devuyst, 2006; Hance et al., 2011). Austin et al. (1973) analyzed the Burrington Oolite in the Mendips area (misinterpreted as the lateral equivalent of the Caninia Oolite). They recorded typical foraminifers of the V2b in Belgium, such as Pojarkovella nibelis (as Quasiendotyra), Klubonibelia (as Mikhailovella) and Nodosarchaediscus. The latter has been later recorded in the upper part of the early Viséan (e.g., Conil et al., 1991), so it cannot be considered currently as a mid Viséan marker. They also recorded an assemblage with Dainella, which they interpreted to be equivalent to the Caninia Oolite. However, these specimens can be attributed to Vissarionovella tujmasensis and V. donzelli, a genus described from the mid Viséan that extended into the late Viséan (Cózar and Vachard, 2001), although it has been rarely documented later from the early Viséan (Kalvoda et al., 2011; Vachard et al., 2018). Subsequently, the assemblage in the Burrington Oolite

is a typical mid Viséan assemblage, without any early Viséan foraminiferal survival. Conil (1963) documented the occurrence of Uralodiscus and Glomodiscus in the mid Viséan in the Campine basin, although in Conil and Lys (1964) only species with reduced contraforts were described from the mid and late Viséan of the Campine basin (e.g. Glomodiscus nudatus and questionably G. lenitortus), that, as has been mentioned, is a relatively usual case. Later, Kimpe et al. (1978) described typical early Viséan foraminifers in the mid and late Viséan of the Brabant Massif in Belgium. They recorded Tournayellidae, Forschiinae, cf. Eotextularia diversa and cf. Dainella sp. in the Argenteau Castle and Glomodiscus miloni in the Railway cutting section West of Bernau. The former assemblage was recorded in outcrops dated as late Viséan, and the latter in rocks dated as mid Viséan (using rugose corals). These authors attributed that fauna to reworking, a process rather common in rocks of the Viséan succession in that region. They did not publish any horizon with mixed fauna, so this inferred reworking is the more plausible interpretation. In Moravia, data are a little ambiguous. Kalvoda (1983, 1990) stated that survival taxa such as Spinobrunsiina, Spinoendothyra, Dainella, Tuberendothyra, Glomodiscus and Uralodiscus were common in the mid Viséan. Spinobrunsiina is a taxon that did not disappear in the early Viséan, and is commonly recorded up to the late Viséan and early Serpukhovian (Vachard et al., 2016). However, the specimen illustrated (Kalvoda, 1990, pl. 3, fig. 6) has to be considered as Spinolaxina, a genus that generally disappeared at the end of the early Viséan. The specimens identified as Dainella are currently included under the genus Vissarionovella that is commonly recorded from the mid Viséan. The specimen illustrated as Glomodiscus is one of those narrow forms that can be recorded up to the late Viséan (see Cózar, 2000). Unfortunately, Uralodiscus, Tuberendothyra and Spinoendothyra were not illustrated. Later, Kalvoda (2002) included the same zonal scheme, but in the stratigraphic range (Kalvoda, 2002, fig. 23) these taxa only crossed up to the base of the mid Viséan or Pojarkovella nibelis Zone, not through the mid Viséan as interpreted in previous studies. This distribution could be compared to those in Morocco in the Skoura boutonnière and in the Russian platform, even though a well-developed, longer persistence of the taxa is not observed. In the southern Montagne Noire in France, Vachard et al. (2018) recorded common species of Eoparastaffella up to the mid Viséan, as well as Brenckleites, Endolaxina, Glomodiscus sp. and Uralodiscus rotundus up to the late Viséan. Some of the specimens were reworked, although some other specimens show good preservation and did not show traces of reworking. In some cases, all the specimens recorded were interpreted as reworked, such as Brenckleites, Dainellidae, Spinolaxina and Florenella up to the Cf6␥. Devuyst et al. (2003) mentioned the occurrence of Uralodiscus in the same bed with Pojarkovella nibelis in the Pengchong section (China), and dainellids in younger beds, interpreting them as survival taxa. Later, Devuyst (2006) published the full list of foraminifers of the Pengchong section, and, above or at the same bed with Pojarkovella nibelis, he listed Endolaxina laxa (as Laxoendothyra), Uralodiscus sp., cf. Conilidiscus sp., Glo-

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modiscus sp. and some species of Eoparastaffella. Other sections in China, such as the Yajiao section, only contain Uralodiscus sp., just in the same bed as the first occurrence of P. nibelis. Hance et al. (2011) revised the Pengchong and Yajiao sections, after questioning some of the specimens previously mentioned, including Ammarchaediscus or Viseidiscus (for taxa previously identified as Uralodiscus), they illustrated Glomodicus (but generally with rather reduced contraforts) and Uralodiscus cf. elongatus. Similar to Moravia, those forms extend only into the basal part of the mid Viséan, but not throughout this substage. Taking into consideration the above described data, it can be summarized that some typical early foraminifers survived into the basal part of the mid Viséan in Moravia and China (previously well-known from the Russian platforms). In contrast, these faunas seem to extend widely into the mid Viséan and parts of the late Viséan in the Western Meseta of Morocco and Montagne Noire in France, although in the latter, the total number of species and genera that survived is not as significant as in the Western Meseta.

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Fig. 5. Palaeogeographical sketch of the Western Palaeotethys for the early Viséan. Data for the reconstruction are mostly based on Simancas et al. (2005) and modifications of Michard et al. (2010). Black areas are the only well-known ophiolites, and the dark areas were emerged lands. Abbreviations: ALZ – AsturLeonese Zone, CZ – Cantabrian Zone, SIF – South Iberia Fault, SMF – South Meseta Fault.

6. Discussion In total, there is a large list of taxa that are recorded much later than in their counterparts in Western Europe: Archaediscus at involutus stage, Brenckleites fragilis, Conilidiscus sp., C. cf. bucculentus, C. settlensis, Cribranopsis foeda, Endolaxina sp., E. laxa, Eoparastaffella macdermoti, Glomodiscus sp., G. lenitortus, G. miloni, G. oblongus, G. praeconvexus, Lapparentidiscus sp., L. bokanensis, L. nanus, Latiendothyranopis aff. menneri solida, Pseudolituotubella sp., P. tenuissima, Pseudoplanoendothyra sp., Septabrunsiina minuta elegantula, Spinobrunsiina sp., Uralodiscus sp., U. elongatus and U. rotundus. They commonly extended their stratigraphic range into the mid Viséan and more rarely into the late Viséan, in the NW Fourhal outcrops and in the north of the Azrou-Khenifra basin. The basins and outcrops revised previously, in neighbouring areas, as well as in the rest of the World, are all developed in distinct tectonic and sedimentological contexts, so it seems not to be plausible that there was a direct relationship between abiotic factors and the survival of the early Viséan foraminifers. In particular, the absence of these survivals into the early Viséan fauna in the surrounding regions does not support the idea that their appearance could be attributed to dispersal from a neighbouring basin (Fig. 5). In terms of palaeolatitude, most of the basins were located in the equatorial belt, with, in some cases, slightly low values of southern latitude. The Montagne Noire and the Western Meseta (both areas where the survivals are more common) were likely situated at a similar latitude (e.g., Torsvik and Cocks, 2004; Stampfli et al., 2013; Izart et al., 2017; Davydov and Cózar, in press). However, both regions were situated in different palaeogeographical settings. The Montagne Noire was located in the southern branch of the Variscan collision (east of the Cantabrian Zone, CZ in Fig. 5), well-connected to the open sea, the Palaeotethys Ocean. In contrast, the Western Meseta was originally located in the northern branch of the Variscan collision, close to the Rheic Ocean Gateway (Davydov and Cózar, in

press, fig. 1A), with a progressive displacement towards Gondwana during the orogeny, with several intervening emerged land masses (Fig. 5). A direct interchange of fauna between the Montagne Noire and the Western Meseta did not seem to exist, and the fauna recorded there was interpreted to be derived from the Laurasian margin, with dispersal through outcrops in the Ossa-Morena Zona of SW Spain (Somerville et al., 2013). The position of the Western Meseta in this corridor for the Mississippian, with some emerged terranes due to the collision, modified the ocean currents for the region, a fact that might explain satisfactorily the late arrival of the mid foraminiferal assemblages to the region, but, by itself, cannot justify its longer preservation. A general isolation of the Western Meseta due to oceanic currents is hard to envisage because later, during the largest late Viséan transgression affecting the Western Palaeotethys, the benthic fauna of the Western Meseta was rather homogenized with that of its counterparts in Western Europe. Foraminiferal markers and assemblages are rather similar to those recorded in surrounding regions. However, some of the surrounding terranes were certainly emerged for most of the early and mid Viséan, as the Coastal Block, Eastern Meseta and Sehoul Zone (Hoepffner et al., 2005), and thus, marine communication for the Western Meseta was only plausible from the West or East (Fig. 5). Furthermore, the Western Meseta is not particularly characterized by many phenomena of speciation, which could justify the total isolation by geographic barriers, and generation of allopatric species. A possible abiotic factor generating a partial isolation is the existence of nearshore currents, generating different temperature gradients to the ocean mass (Schopf, 1979). These nearshore currents would flow predominantly in the northeastern part of the present-day Western Meseta, where this effect of longer persistence for the foraminifers is sharpest. Another important factor that would have favoured the longtime persistence of the foraminifers could be the late arrival of the typical mid Viséan assemblages (Vachard and Tahiri, 1991)

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Fig. 6. Main ecological patterns of distribution of the Viséan foraminifers; the resulting assemblage is observed in the grey area. (A) Early and mid Viséan stocks share their stratigraphic and geographic ranges during the mid Viséan, whereas the early Viséan stock is not recorded during the late Viséan (Skoura, NE Sidi Bettache, NE Oulmès-El Hamman). (B) Early Viséan stock shares its stratigraphic and geographic ranges with the foraminiferal stocks for the mid and late Viséan (NW Fourhal). (C) Early Viséan stock overlapped its stratigraphic ranges but not its biogeographic limits through the mid and late Viséan (north of Azrou-Khenifra). Note that the mid and late Viséan foraminiferal stocks always overlapped in time and geographically.

and subsequent lack of biotic competition or species interactions for the same ecological niches (Anderson et al., 2002). These latter authors proposed a model for large-scale biogeographic areas based mostly on biotic interaction between two species. Usually, these patterns of competition or species interaction are better demonstrated in small-scale biogeographic differentiation (Wiens, 2011), as can be considered for the Western Meseta, which is of a regional scale, and not affecting large areas (as could the entire Western Palaeotethys). In most of the Western Palaeotethys basins, the arrival of the mid Viséan assemblages led to the disappearance of a suite of typical early Viséan foraminifers, which can be interpreted as competition for the same ecological niches. Peculiarities in the Western Meseta show a few regions where: (1) early and mid Viséan faunas overlapped their stratigraphic and geographic ranges up to the mid Viséan (Skoura, NE Sidi Bettache, NE Oulmès-El Hamman), whereas during the late Viséan a mixture of mid and late Viséan foraminifers are recorded (Fig. 6A); (2) early Viséan faunas overlapped its stratigraphic and geographic ranges during the mid Viséan with typical foraminifers of this substage, and finally, during the late Viséan, foraminifers of the three substages occurred (NW Fourhal) (Fig. 6B); and (3), there are areas where faunas overlapped their stratigraphic ranges but not their biogeographic limits through the mid and late Viséan (north of Azrou-Khenifra), being recorded in separated stratigraphic levels (Fig. 6C). The three cases are compelling cases where the taxa show geographic range overlap and indicate the dispersal is limited by species interactions (Wiens, 2011). In particular, in sections of the northern Azrou-Khenifra basin, there is competitive release, allowing the early Viséan foraminifers to expand their geographic ranges (as well as their stratigraphic ranges) in the absence of the mid and late Viséan foraminifers, establishing species limits for a given area and a given time slice (Fig. 6C). The palaeontological data from the Western Meseta combine historical biogeography and trait evolution with time-calibrated phylogenies, and thus can contribute to the study of the Eltonian niche (Wiens, 2011). In modern ecological studies, the lack of

historical records may have lost their perspective if their biogeographic ranges are a matter of biogeographic provinces or local biogeographic effects. The combination of both factors is rarely considered (Moen et al., 2009). However, it is not clear why, in some sections, early Viséan assemblages co-habited, sharing space and time with the newly arrived species of the mid or late Viséan. Similar differences are observed in modern ecological patterns, with no obvious explanation, suggesting that there are still many questions to be studied about geographic patterns (Wiens, 2011). In modern ecology, two hypotheses that could explain the differences between the regions of the Western Meseta can be found: ecological limits in the number of species by regions over time (Rabosky, 2009) or niches saturated in ecological similar species over time (Loreau, 2000). However, all these factors are generally associated also with species interactions. The most plausible explanation is that the survival of the early Viséan foraminiferal assemblages in the Western Meseta of Morocco is a matter of species interaction and competition, a fact primarily triggered by the late arrival of the younger foraminiferal assemblages in the region. 7. Conclusions Taxonomic, sedimentological and taphonomical analyses of typical early Viséan foraminiferal assemblages recorded in middle and upper Viséan stratigraphic sections of the Western Meseta of Morocco suggest that they are composed mostly of autochthonous and para-autochthonous assemblages with a minor percentage of reworked specimens, although, locally, important percentages of reworking occur. Thus, most of these faunas readily crossed the middle Viséan boundary, where most of them were extinct in the Palaeotethys. Analysis of the stratigraphical occurrence of the early Viséan assemblages demonstrates that there are three basic patterns: early and mid Viséan assemblages sharing similar stratigraphic and biogeographic ranges (Skoura, North Sidi Bettache, North Oulmès and El Hammam Ridge); early Viséan forms shar-

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ing the same stratigraphic and biogeographic ranges with mid and late Viséan foraminiferal assemblages (NW Fourhal); and early Viséan assemblages sharing stratigraphic ranges but not biogeographic ranges with mid and late Viséan foraminiferal assemblages (north of Azrou-Khenifra). Revision of the surrounding Viséan basins does not show survival of the early Viséan foraminiferal assemblages apart from some rare species, so no similar patterns are recognized. A worldwide comparison only suggests that the early Viséan foraminifers persisted in the lowermost part of the mid Viséan (but not through this substage) in Moravia and China, whereas only in southern France (Montagne Noire), typical early Viséan foraminifers survived widely up to the late Viséan, although in less abundance and diversity than in the Western Meseta. These incongruous results allow the most important abiotic factors which triggered the survival of the fauna (e.g., tectonic setting, environmental setting, isolation by oceanic currents or land barriers, palaeotemperature, palaeolatitude) to be discarded. Nevertheless, some local abiotic factors could have favoured the development of these patterns (e.g., nearshore currents), but the persistence of the early Viséan foraminifers in much younger rocks is mostly interpreted as due to species interaction and competition, which could be primarily triggered by the late arrival of the mid Viséan foraminifers to the Western Meseta.

Acknowledgements We would like to thank S. Arefifard and Qing-Yi Sheng by their constructive comments, and the editorial work by Yi-Chun Zhang. The authors thank Spencer Lucas for reviewing the earlier draft of the manuscript and English improvement. Fieldwork for PC and IC was funded by the Spanish Ministerio de Ciencia, Innovación y Universidades (project CGL2016-78738BTE).

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