Biostratigraphy and paleobiogeographic affinities of the Jurassic brachiopod assemblages from Sierra Espuña ( Maláguide Complex , Internal Betic Zones , Spain )

The assemblages of Early Jurassic brachiopods (Pliensbachian Toarcian) from Sierra Espuña (Murcia Province, SE Spain) are described. This is the only area in the Internal Zones of the Betic Cordillera, corresponding to the margins of the Alborán Terrane, where Jurassic brachiopods are known to occur. In the tectonic Unit of Morrón de Totana (more southward located) assemblage MT1 of Late Pliensbachian age has been characterized. This assemblage has been subdivided into three successive sub-assemblages: MT1a (Algovianum Zone), MT1b (Emaciatum Zone, Solare Subzone) and MT1c (Emaciatum Zone, Elisa Subzone). Northward, in the Perona tectonic Unit two distinct assemblages, P1 (Latest Sinemurian Early Pliensbachian) and P2 (Early Toarcian, Serpentinum Zone) have been recognized. Differences between the assemblages from the two tectonic units are evident after the paleobiogeographical analysis. In the Morrón de Totana Unit, taxa with Mediterranean affinities occur. MT1 assemblage is very similar to assemblages previously known in the Eastern Subbetic as well as in other areas of the Mediterranean Province. In the Perona Unit the Mediterranean affinity of the assemblages is not so evident. P1 Assemblage consists of widely distributed taxa, lacking in the most characteristic elements of the Mediterranean Province which, however, are present in neighbouring Betic areas. P2 Assemblage belongs to the Spanish Province that develops in Western Tethys after the Early Toarcian Mass Extinction Event. The occurrence in this assemblage of Prionorhynchia aff. msougari Rousselle, until now only found in North Africa, indicates a closer connection of the Perona Unit with the African paleomargin of the Tethys than with the South Iberian paleomargin. The paleobiogeographical data suggest a more southern and marginal (close to epicontinental areas) position of the Perona Unit than the Morrón de Totana Unit.


Introduction
The Early-Middle Jurassic transition paleogeography of the westernmost Tethys area was characterized by two major plates separated by the Central Atlantic Ocean in a Red Sea-type stage.The northern plate one was the Laurasian and the southern one was the African Plate.The Tethys Ocean was widening to the East.The Iberian Subplate was located in the southern margin of the northern plate, structuring the South Iberian margin (Martín-Algarra and Vera, 2004), which originated the External Betic Zone during the Alpine Orogenesis, and the northwestern margin of the African plate that evolved into the External Rif Zone.In the Tethys Ocean, the AlKaPeCa Microplate (Alboran, Kabilyan, Peloritanian and Calabria;Bouillin et al., 1986), also known as Mesomediterranean plate (Martín-Algarra et al., 1992;Guerrera et al., 1993), comprises the internal zones of the Betic-Rif, Tellian, Kabylian, Calabria-Peloritanian and Southern Apennine Chains.The paleolatitude of the South Iberian paleomargin was about 30º N, whereas the AlKaPeCa Microplate was probably situated at a similar paleolatitude, but more eastward (Ziegler, 1990;Bassoullet et al., 1993;Vera, 2001;Stampfli and Borel, 2004).The Tertiary tectogenesis divided this microplate in different terranes separated by the extension of the Western Mediterranean Sea: Alboran, Kabilyan, Peloritanian, and Calabria.
The Alboran Terrane comprises the internal zones of the Betic and Rif Cordilleras.It is built up by the stacking of three complexes, called from bottom to top: Nevadofilabride, Alpujarride-Sebtide and Malaguide-Ghomaride (Sanz de Galdeano, 1997).The Nevadofilabride and Alpujarride units are composed mainly of Paleozoic and Tri-assic metamorphic rocks, while the Malaguide Complex includes very low grade Paleozoic metamorphic rocks and a Meso-Cenozoic sedimentary cover.In general, the scarcity of Jurassic non-metamorphosed successions, and the intense poli-phase tectonics have hampered a detailed description of the geodynamic evolution of the Jurassic from the internal zones.
The Malaguide Complex outcropped mainly from the Málaga Province (westward) to the Murcia Province (eastward).Analogous sediments to the Malaguide Complex are also present in the northern Rif Cordillera (Morocco), where they are known as the Ghomaride Complex (Maaté, 1996).The Sierra Espuña area, in the Murcia Province (southern Spain), is probably the most extensive, less deformed and best exposed Jurassic-Cretaceous outcrop belonging to the Malaguide-Ghomaride complexes in the Internal Betic-Rif Chain (Fig. 1A).
Very few studies have been focused in the Mesozoic paleontological content of the internal zones of the AlKa-PeCa Microplate due to the lack of non-metamorphic outcrops, as the Ghomaride (Maaté, 1996), or the Tell (Alméras et al., 2007).Within the Malaguide Complex, the extensive and continuous Jurassic outcrops containing macroinvertebrate fossils in Sierra Espuña are noteworthy, which give the possibility to study the ammonite fauna, providing a precise biostratigraphic framework for the Jurassic (Fallot, 1945;Peyre and Peyre, 1960;Paquet, 1969;Geyer andHinkelbein, 1974 andCaracuel et al., 2006).
Giving continuity to the research by Caracuel et al. (2006) the aim of this paper is to study the brachiopod assemblages in the Malaguide Complex, focusing on the Early-Middle Jurassic transition outcropping in the Si-
The first work in which brachiopod species are identified in Sierra Espuña is due to Paquet (1969).This author assigned to the Middle Domerian a level of red limestones with ferruginous oolites in the section of Morrón de Totana, on the basis of an abundant ammonite fauna found together with Spiriferina cf.cantianensis?Canavari, Spiriferina sp.aff.decipiens Böse and Schlosser, S. appenninica Canavari in Haas, S. falloti Corroy, S. cf.gibba Seguenza, Rhynchonella cf.persinuata Rau, R. cf.restricta Parona, R. quadrata Buckman, R. sp., Terebratula sphenoidalis Meneghini and Zeilleria sp.In the Perona area this same author (Paquet, op. cit.) reported the presence of Spiriferina ascendens Deslongchamps, S. gr.haueri Suess, S. falloti?Corroy, Rhynchonella curviceps Quenstedt, R. sp., Zeilleria roemeri Schlotheim and Z. cf.roemeri in red ferruginous limestones assigned to the Early Pliensbachian.erra Espuña area (Fig. 1B).The similarity of many of the studied taxa with those recorded in the Eastern Subbetic and other areas of the Western Tethys will also provide new criteria aiding to place more precisely these associations within the established chronostratigraphy as well as their paleobiogeographical affinities.

Materials and Methods
Brachiopods were collected in three stratigraphic sections which are little-tectonized and contain the most complete and fossil-rich successions: Tres Carrascas, Malvariche, and Perona (Fig. 1).The taxa reported by previous authors from the lower part of the Perona section and from Prat Mayor (Paquet, 1969;Geyer and Hinkelbein, 1974) have been also considered in this study, after an up-date of their taxonomic determinations.Taphonomic observations were considered in the Malvariche section, where the taxonomical composition of the samples is clearly biased.
The ammonite zonal/subzonal scheme follows Cariou and Hantzpergue (1997) for the Pliensbachian and Toarcian of the Mediterranean Domain.Chronostratigraphic data from Caracuel et al. (2006) in Sierra Espuña area were also considered.Facies characteristics of the beds containing the studied brachiopods were described and related to the regional stratigraphic framework.Twelve thin sections were also used for characterizing some microfacies and textures, especially from the ferruginous oolithic limestone interval.For paleobiogeographical interpretations, species representation, species richness and comparison of shared species with other basins of the Western Tethys were considered.

Geological setting and facies distribution
The Malaguide units outcropping in the Sierra Espuña area bounds tectonically with the Alpujarride Complex to the SE and with the External Betic Zones (Subbetic) to the NW (Fig. 1B).
The Early-Middle Jurassic transition sediments of Morrón de Totana Unit outcrop with lateral continuity for more than 10 km over a Paleozoic to Triassic succession, from the Prat Mayor area to the Malvariche Valley area.The synthetic stratigraphic succession distinguished in this unit it is shown in Fig. 3.
The lowermost rocks of Jurassic age in this succession are of dolomitic facies traditionally attributed to the Earliest Jurassic (probably Hettangian).Overlying them a thick set of oolitic-pisolitic white limestones evolving upwards to pinkish oncolitic-rodolitic limestones are present.These are sometimes massive frequently crossbedded and sometimes brecciated beds, often with thickness of ten metres.Texturally they are grainstones to packstones (occasionally rudstones).The microfacies of this portion of the section was analysed by Caracuel et al. (2006), showing algal remains, ostracods, sponges and benthic foraminifera.Kampschuur et al. (1972)  a Sinemurian to Pliensbachian age for these limestones.
Overlying this sequence red-pinkish crinoidal grainstones are present, with intercalations in the upper part, of sandy limestones first and ferruginous oo-oncolitic limestone towards the top.The brachiopods studied in this work from Malvariche and Tres Carrascas Sections came from these sandy and crinoidal grainstones levels, characterized by a rich ammonite content indicating a Domerian to Toarcian age.The oolites/oncoids consist of concentric growth of Fe-Mn laminae around a nucleus (Fig. 2e-f).Finally, in the Middle Jurassic well-bedded micritic (oolitic in the basal levels) and crinoidal limestones facies with abundant cherty nodules prevail.
In the Perona Unit, the basal sequences of the Early Jurassic carbonate succession show generally the same facies found in the Morrón de Totana Unit: dolomitic at the base, followed by oolitic white limestones and red crinoidal limestones.However, over the crinoidal tract a set of red, ferruginous detritic limestones appear, fol-

Species
lowed by an alternation of yellowish marls and marly/ silty limestones levels with brachiopods, some ammonoids and trace-fossils of Thalassinoides type.

Preservation of the brachiopod shells.
A total number of 438 brachiopods were studied.The detailed provenance of the brachiopods is indicated in Table 1.Several specimens of the more representative taxa are shown in figures 4 and 5.In most localities the brachiopods are well preserved, with nearly 80% of fossils preserving the shell and showing a low degree of breakage.However, a low taxonomic diversity, high breakage and reworking is found in the Malvariche section, implying a preferential accumulation of flat and subcircular-elliptic valves such as the dorsal valves of spiriferinids and similarly shaped pectinids; the ratio of dorsal/ventral valves Table 1.-Detailed ocurrence of Early Jurassic brachiopod species in the Sierra Espuña studied sections and their stratigraphical distribution in the Western Tethys.The stratigraphic data were obtained from the references indicated in section 5 in the text; additional data from Alméras (1964).Tabla 1.-Especies de braquiópodos del Jurásico Inferior registrados en los afloramientos estudiados en Sierra Espuña y su distribución estratigráfica en el Tethys Occidental.El origen de los datos de las distribuciones estratigráficas se cita en el texto, completándose con las citas de Alméras (1964).
sampled is 78.95% / 21.95%.In this locality, ammonoids show neomorphosed phosphated shells and sometimes with Fe-Mn oxide crusts.Internal recrystallization of the specimens is common though not dominant in all the studied assemblages.Occasionally, juvenile specimens with ferruginous crusts are found in the Fe-oolites intervals.In the Perona section, Soaresirhynchia bouchardi specimens frequently display fosildiagenetic deformations and tectonically compressed specimens.
In most samples no size sorting was observed but occasionally some mono-or di-specific assemblages were found in Malvariche and in Perona.In the first case it is clearly a consequence of reworking, whereas in the second case the assemblages, highly dominated by Soaresirhynchia bouchardi, seem to be related to unique colonization and opportunism strategies of this species (García Joral and Goy, 2000, Gahr, 2005, Graziano et al., 2006, García Joral et al., 2011).
As in the previous bed, some brachiopod taxa with wide stratigraphic ranges such as Prionorhynchia polyptycha, Lobothyris punctata, Liospiriferina rostrata, L. alpina, L. gryphoidea or C. tumida, are found in this level together with other taxa restricted to the Late Pliensbachian.These are Prionorhynchia quinqueplicata, Prionorhynchia gignouxi, recorded in the Subbetic in the Late Pliensbachian (Alméras et al., 1993;Baeza-Carratalá, 2008), and Rhapidothyris delorenzoi, recorded in the Spinatum Zone of the Bakony Mountains (Hungary) by Vörös (2009), and between the Lavinianum and the lowermost Polymorphum (Early Toarcian) Zones in the Subbetic of Alicante and North Murcia by Baeza-Carratalá (2010).

Malvariche Section
The brachiopods studied in this section were collected in yellowish sandy limestones with abundant iron oxides and phosphates.They were found together with numerous pectinids and some ammonites and belemnites.All brachiopods were identified as belonging to the genus Liospiriferina (83% Liospiriferina alpina + 17% Liospiriferina cf.rostrata), both species presenting a wide stratigraphic range.According to Caracuel et al. (2006), the ammonite fauna in these levels is abundant, with taxa of Late Pliensbachian age (Algovianum Zone, Ragazzoni, Bertrandi and Accuratum Subzones) found together with others of a somewhat older age (Lavinianum Zone): Fuciniceras cornacaldense (Tausch), Fieldingiceras fieldingii (Reynes), and Becheiceras bechei (Sowerby).

Prat Mayor Section
In the locality of Prat Mayor, Paquet (1969)  Later, Geyer and Hinkelbein (1974) found Rhynchonella wilfridi Jiménez de Cisneros, 1923 and Terebratulidae sp.indet. in grey to yellowish marls and limestones of the same locality.Rhynchonella wilfridi has been considered as synonymous of Prionorhynchia quinqueplicata (Zieten) by Alméras et al. (1993), Baeza-Carratalá et al. (2004) and Baeza-Carratalá (2008).As indicated above, this species has a stratigraphic range restricted to the Upper Pliensbachian in the Eastern Subbetic and other areas of Western Tethys.This age is supported by the ammonite fauna recorded by Geyer and Hinkelbein (op.cit.) composed by the genera Arieticeras/Canavaria/Fontanelliceras, Lioceratoides and Catacoeloceras.
The same authors (Geyer and Hinkelbein, 1974) in a nearby locality within the Morrón de Totana Unit, reported a tract with brachiopods stratigraphically above the previously described, consisting also of grey to yellowish marls and limestones but partially nodular and with ferruginous oolites.They identified "Rhynchonella" ssp., Spiriferina alpina (=Liospiriferina alpina) and Spiriferina angulata (=Cisnerospira angulata), together with an ammonite association comprising Lytoceras sp.ex.gr.sepositum Meneghini, Peronoceras cf.millarense Monestier, Catacoeloceras tethysi Géczy, Arieticeras sp., Hildoceras cf.bifrons (Bruguière), and H. cf.graecum Renz.In this assemblage species from the Early and Middle Toarcian appear, hence the authors admitted a certain condensation in these levels.This condensation agrees with the brachiopod data, since Liospiriferina alpina and Cisnerospira angulata, though ranging until the earliest Toarcian in some areas, do not surpass the Early Toarcian Mass Extinction Event at the base of the Serpentinum (=Falciferum) Zone.This event marks the final extinction of the order Spiriferinida (cf.Comas-Rengifo et al., 2006;García Joral et al., 2011).The co-occurrences of these species with H. bifrons from the Middle Toarcian is thus likely related to temporal averaging of Early and Middle Toarcian specimens.

Perona Section
The brachiopods were collected in two beds (Pe-1 and Pe-2), within a tract of yellowish marly limestones that alternates with silty limestones, occasionally slightly nodular.In both beds, the assemblages are clearly dominated by Soaresirhynchia bouchardi (Davidson), with a few accompanying species.
The specimens of Soaresirhynchia bouchardi collected in this locality show high morphological variability displayed elsewhere in neighbouring basins, so that it is possible to recognize the distinct morphotypes (calva, penichensis, flamandi) that were described by several authors and sometimes considered to represent different species (Dubar, 1931;Rousselle, 1978;Alméras, 1994, Alméras andFauré, 2000;Alméras et al., 2007Alméras et al., , 2010 among others).However, no boundaries between the morphogroups are recognizable and the collected specimens have been assigned to a single species, following the same criteria considered in Comas-Rengifo andGoy (1975), García Joral andGoy (2000) or García Joral et al. (2011).
Other brachiopod taxa identified in Pe-1 bed are: Pseudogibbirhynchia sp.(a single large-sized specimen with the bifurcated rounded ribs typical of this genus but not assignable to any known species), Homoeorhynchia sp., Telothyris jauberti (Deslongchamps), and T. pyrenaica (Dubar).
These brachiopods were collected in the same levels where Caracuel et al. (2006) identified Dactylioceras (Eodactylites) sp. and Hildaites striatus Guex, that characterize the Polymorphum and Serpentinum Zones of the Early Toarcian.An age restricted to the Serpentinum Zone can now be proposed for these levels, since the brachiopod assemblages found in Pe-1 and Pe-2 beds never occur below the Serpentinum Zone in the neighbouring basins.Soaresirhynchia bouchardi characterizes the El-egantulum Subzone of the Serpentinum Zone in the Iberian Range (García Joral and Goy, 2000), having a similar range in other basins, from southern England to Morocco and from Portugal to the Balkans (Ager, 1962;Rousselle, 1973;Alméras, 1994;Gakovic and Tchoumatchenco, 1994;Alméras and Fauré, 2000;Graziano et al., 2006;Alméras et al., 2007Alméras et al., , 2010)).This species has been recognized in a similar stratigraphic position in other areas of the Betic Range, such as Mallorca Island (Álvaro et al., 1989), or the eastern Subbetic (Baeza-Carratalá, 2008).
Pseudogibbirhynchia jurensis and Telothyris jauberti are recorded in the Iberian Range in the Serpentinum Zone, and T. pyrenaica is recorded both in the Serpentinum and in the Bifrons Zones (García Joral and Goy, 2000).A similar distribution is observed for these taxa in other regions of Western Tethys (cf.Alméras et al., 2007Alméras et al., , 2010;;García Joral et al., 2011 for a revision).Prionorhynchia msougari is recorded in the Middle-Upper Toarcian of the High Atlas (Rousselle, 1973;Alméras et al., 1993) and from the Lower Toarcian of Western Algeria, where it is found with species of the genera Soaresirhynchia and Telothyris (Alméras et al., 2007).

Brachiopod Assemblages
Brachiopod assemblages from the Morrón de Totana and Perona Units can be subdivided into three major groups with temporal and paleobiogeographical significance: MT1 Assemblage: It occurs in the Morrón de Totana Unit and is recorded in the Tres Carrascas, Malvariche and Prat Mayor Sections.It is dominated by large-sized Prionorhynchia specimens, and by smooth-shelled spiriferinids.Intraplicate terebratulids and axiniform zeillerids are scarce (Fig. 4).The taxa represented in this assemblage are characteristic of the Late Pliensbachian of the Mediterranean Province.An analogous assemblage has been reported in a similar stratigraphic position in the Eastern Subbetic (Baeza-Carratalá, 2008).

Contributions to Paleogeography and Paleobiogeography
The paleogeographical context for the Jurassic of Sierra Espuña has been thoroughly studied (e.g.: Martín-Martín, 1996;Martín-Martín et al., 2006;Caracuel et al., 2006).At the beginning of the Jurassic, this area was located near the southern margin of the AlKaPeCa Microplate From a paleobiogeographical point of view, this area was situated near the margins of Africa and Iberia (Martín-Algarra and Vera, 2004), close to the western boundary between the Mediterranean and the European Provinces (the latter also referred to as the Northwestern European) (cf.Ager, 1967Ager, , 1971Ager, , 1973;;Vörös, 1977Vörös, , 1984Vörös, , 2002Vörös, , 2005;;Manceñido, 2002).(Alboran, Kabilyan, Peloritanian, and Calabria;Bouillin et al., 1986) as part of a shallow and restricted platform.This platform broke-up during the Early Jurassic, leading to an open shelf depositional environment with variable depth and a complex bottom due to block faulting and tilting (Martín-Martín et al., 2006).Table 2. Comparison of shared species in the brachiopod assemblages recognized in the Sierra Espuña and other representative areas of Western Tethys, in total number and percentages.Tabla 2. Número y porcentaje de especies de las asociaciones de braquiópodos reconocidas en Sierra Espuña presentes en otras áreas representativas del Tethys occidental.

Assemblage
Eastern Subbetic (Baeza-Carratalá, 2008) Bakony (Vörös, 2009) Western Algeria (Alméras et al., 2007) Iberian Range (Comas-Rengifo, 1992;García Joral and Goy, 2000) Pyrennees (Alméras and Fauré, 2000) MT1 ( NT=17  The brachiopod assemblages described in this paper indicate that the two tectonic Units of the Malaguide complex differed in their paleobiogeographic affinity. In the Morrón de Totana Unit, assemblage MT1 has clear Mediterranean affinities, with a high percentage of species (82%) also found in the eastern Subbetic and in other typical Mediterranean domains as Bakony Moun-tains (47%), whereas this affinity is low (23,5%) when compared with European domains (Iberian Range and Pyrenees) and with the African margin (Table 2).Moreover, three of the four species shared with European regions (Liospiriferina alpina, L. rostrata and Lobothyris punctata) can be considered ubiquitous, and are also found in the Mediterranean Province.The fourth, Cal-lospiriferina tumida, considered properly European, has been however also reported in the eastern Subbetic (Baeza-Carratalá, 2008).
The two assemblages described in the Perona Unit are not so clearly related to the Mediterranean Province (Table 2).P1 is composed by widely distributed species.The most representative taxa are Liospiriferina rostrata, Gibbirhynchia curviceps, Zeilleria roemeri, and Cisnerospira aff.adscendens.The three former species are present in the Pyrenees, the Iberian Range, the Betic Ranges and Western Algeria, whereas only the ubiquitous G. curviceps is reported also in the Bakony.Cisnerospira aff.adscendens seems to be endemic to the Betic area.The absence of the typical taxa of the Mediterranean Early Pliensbachian (Apringia, Pisirhynchia, Linguithyris, etc.) or even of the taxa that characterized this interval in the eastern Subbetic (Tauromenia, Lychnothyris, Praesphaeroidothyris, Securina;cf. Baeza-Carratalá, 2011;Baeza-Carratalá and García Joral, in press) suggests a closer relation of this assemblage with those of the European and African margin than with the faunas characteristic of the Mediterranean Province.P1 Assemblage come from red ferruginous detritic limestones, probably deposited in slightly more "epicontinental" conditions than the ferruginous silty limestones with Fe-oolites from which MT1 assemblage comes.
Having in mind that the available paleogeographic data indicate proximity (Martín-Martín, 1996, estimated a distance of 10-12 km between the two tectonic units in the Early Jurassic) and that all the species in P1 assemblage also occurs in the eastern Subbetic (a paleomargin from a different subplate separated at least 500 km from Sierra Espuña; cf.Martín-Algarra and Vera, 2004), the differences in taxonomical composition could be related to paleoenvironmental differences.The Perona Unit probably corresponds to environments more influenced by continental areas than in the Morrón de Totana Unit and in the eastern Subbetic, where thicker, more pelagic and with higher deeping rate succesions are represented.
P2 Assemblage is characteristic of the Spanish Province (Choffat, 1880, Dubar, 1931, Delance, 1972, García Joral and Goy, 1984, 2004) that builds up in the Westernmost Tethys after the provincialism fades as a consequence of the Early Toarcian Mass Extinction Event (ETMEE).The ecological void caused by the ETMEE in both European and Mediterranean areas was filled in by Soaresirhynchia bouchardi in the aftermath of the Extinction, a species with opportunist features (García Joral and Goy, 2000;Gahr, 2005, García Joral et al., 2011), and later on by other species, Homoeorhynchia and Telothyris being the more representative genera.S. bouchardi extends over a wide area of Western Tethys, as stated earlier, whereas the subsequent assemblage is more restricted to westernmost areas, including the Andean Region in South America (Manceñido, 2002).The development of the Spanish Province lasts only until the Aalenian, when the former European -Mediterranean paleobiogeographical pattern is re-established.
The taxa of the Spanish Province have not been described in the core areas of the Mediterranean province, such as in the Southern Alps, Transdanubian Ranges or Sicily.Thus, the Spanish Province can be considered as "marginal" in the Tethyan areas, though the absence of its taxa might also be a consequence of worse preservation conditions.A particularity of P2 Assemblage is the occurrence of Prionorhynchia aff.msougari.This species has not been found in the European domains of the Spanish Province, but they have been reported in Morocco (Rousselle, 1973) and Algeria (Alméras et al., 1993(Alméras et al., , 2007)).This suggests that P2 Assemblage might be more linked to the African margin than to the European one.
The paleobiogeographical and paleoecological differences established between the Morrón de Totana and the Perona Units based on their brachiopod assemblages are in good accordance with the geodynamic structural model of the Sierra Espuña area (Martín-Martín, 1996;Martín-Martín and Martín-Algarra, 1997), in which the Perona Unit was placed not far away, but in a more southern position than the Morrón de Totana Unit.

Conclusions
The Sierra Espuña area has yielded the only Jurassic brachiopod record from the Internal Zones of the Betic Cordillera.Their description increases notably the knowledge of the group in the paleomargin of AlKaPe-Ca microplate, where until now only the Algerian Tell brachiopods had been described.The studied specimens have been assigned to 28 species belonging to 17 genera.Prionorhynchia aff.msougari is identified for the first time in the Iberian Peninsula.
Three assemblages were recognised on the basis of the specimens collected in the two tectonic units defined for this region: the Morrón de Totana Unit (MT1) and the Perona Unit (P1 and P2).MT1 Assemblage has a Late Pliensbachian age and shows clear Mediterranean affinities.P1 Assemblage is of Latest Sinemurian -Early Pliensbachian age, and is mostly constituted by ubiquitous taxa.P2 Assemblage belongs to the Spanish Province that builds up after the Early Toarcian Mass Extinction Event, occupying a wider area of the westernmost Tethys.The occurrence in this assemblage of Priono-la Société des Sciences Naturelles de l'Ouest de la France supplément hors-série: 1-131.Alméras, Y., Ameur, M., Elmi, S. (1993) rhynchia aff.msougari suggests a closer connection of the Perona Unit to the African areas than to the European within the Spanish Province.
The paleobiogeographical affinities of these brachiopod assemblages agrees with the geodynamic stucturing model of the Sierra Espuña area, that implies a relative more southern position for the Perona Unit.From the brachiopod assemblages a more marginal position, close to the epicontinental areas, is inferred for this unit.
recorded Terebratula sp. and Rhynconella wilfridi Jiménez de Cisneros in grey to yellowish limestones from the Late Pliensbachian of Prat Mayor, and Rhynchonella sp., Spiriferina angulata, and S. alpina in weakly nodular limestones and marly limestones from the Early Toarcian of Morrón de Totana. suggest