Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates of...

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Senckenbergiana lethaea 87 (�) �–39 5 text-figs, �� pls Frankfurt am Main, 30.06. 2007

Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates of western Tethys

With 5 Text-figures and �� Plates

Marcelle K. BouDagher-FaDel & Dan W. J. Bosence

A b s t r a c t

Low-latitude carbonate platforms dominated the southern and northern margins and microplates of west-ern Tethys during the Early Jurassic. However, these have proved difficult to date in the past partly due to a lack of study but also a perception of low biotic diversity following the Triassic-Jurassic extinction event. Detailed logging and sampling of seven continuously exposed sections of Lower Jurassic shallow-marine carbonates in Gibraltar, Morocco, Tunisia, Greece, Italy, and in Spain reveals the occurrence of �2 benthic foraminifera species. These taxa are described and illustrated, and include: one new species Textulariopsis sinemurensis, new records for the Early Jurassic, and new records for this region. Five new biozones for the late Hettangian to early Pliensbachian time interval are erected from the consecutive appearance of benthic foraminifera within these sections, from top to base: Lituosepta compressa, Lituosepta recoaren-sis, Everticyclammina praevirguliana, Siphovalvulina colomi and Siphovalvulina gibraltarensis. This has enabled the correlation of these sections for the first time from the northern and southern Tethys margins and from the Apuleian and Pelagonian microplates within Tethys. The foraminiferal occurrences indicate a similarity of associations from the southern and northern shallow carbonate shelves of Tethys, whilst re-flecting the relative isolation of the Apuleian and Pelagonian microplates within Tethys. The foraminifera show a progressive diversification of forms through the late Hettangian to early Pliensbachian interval that is interpreted to reflect the evolutionary recovery and diversification of these biotas following the Triassic-Jurassic extinction event.

K e y w o r d s : Benthic foraminifera, biozones, palaeoenvironment, Jurassic, western Tethys.

Dr. Marcelle BouDagher-FaDel, Department of Earth Sciences, University College London, Gower Street, LONDON, WC�E 6BT, UK. Prof. Dan W. J. Bosence, Department of Geology, Royal Holloway University of London, Egham, SURREY, TW20 0EX, UK.

Introduction

The mass extinction in the marine realm at the end Triassic affected many groups of benthic foraminifera. However, a few survivors continued into the Jurassic but these gradu-ally disappeared towards the Cretaceous (Basov & Kuznetsova 2000). Of the different suborders of foraminifera present in the Jurassic, the agglutinated foraminifera were the only group that witnessed significant development in the Tethys Ocean, with taxa becoming large, complicated, and forming many lineages. The Early Jurassic witnessed the steady evolution of

textulariids from small, simple forms to internally complicated forms that became abundant from the Pliensbachian onwards. These later forms have particular morphological features that are recognisable throughout the shallow-marine facies of Early Jurassic carbonate platforms of Tethys. Simple forms of Siphovalvulina spp. and Textularia dominated the early Sine-murian (BouDagher-FaDel et al. 200�). The mid to late Sine-murian has forms with internal pillars, such as Pseudopfende-rina, and fine alveoles in the walls, such as Everticyclammina; both genera were prominent in the Western Tethys or Mesogea (Fugagnoli �996). Such alveoles developed into larger and

BouDagher-FaDel & Bosence: Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates...2

more consistent alveoles in taxa in the Pliensbachian, and in the Late Jurassic and Cretaceous (septFontaine �98�). Such foraminifera became very important components of Mesozoic rocks.

The Jurassic larger benthic foraminifera have been system-atically studied on a regional scale by a number of authors. Early Jurassic Hettangian-Sinemurian foraminifera from the present-day Mediterranean region were studied by septFon-taine (�98�), Fugagnoli (2000; 2004), and by nouJaiM clarK and BouDagher-FaDel (2003; 2004) who proposed standard Jurassic biozonations for the Mesogean realm on the basis of foraminiferal generic ranges and assemblages. The early Sinumerian is characterized by siphovalvulines, and more complex and diverse imperforate foraminifera appeared only in the late Sinemurian and early Pliensbachian in Tethys (höt-tinger �967; septFontaine �984). This appears to hold true for

the Tethyan realm in general (cf. sartorio & venturini �988); for the extensive carbonate platform (garcía-hernánDez et al. �978; vera �988; rey �997, p. 235; BouDagher-FaDel et al. 200�) that now forms part of the External Zones of the Betic Cordillera of southern Spain (cf. gonzález-Donoso et al. �974; Braga et al. �98�), and for the Lower Jurassic (Calcare Massiccio) of Italy; in the north (Bosellini & Broglio loriga �97�; castellarin �972, Fugagnoli & Broglio loriga �998), the south (sartoni & crescenti �962) and the central Apen-nine areas (e.g. chiocchini et al., �994, Barattolo & Bigozzi �996). cariou & hantzpergue (�997) coordinated research by the “Groupe Français d’Étude du Jurassique” that improved stratigraphical calibration of many fossil groups including those of the larger benthic foraminifera of western Europe and the Mediterranean. This paper describes benthic foraminifera, algae, and a cyanobacterium that form a distinctive Early Jurassic biota within peritidal carbonate platform environ-

Fig. �. Locations of logged sections

BouDagher-FaDel & Bosence: Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates... 3

ments in western Tethys. A new species of Textulariopsis is de-scribed and this constitutes the first record of this taxon from the western Tethys region, and confirms that this seemingly largely Cretaceous genus had originated by the Early Jurassic.

The area studied in this work includes the southern (Mo-rocco and Tunisia) and northern (Spain, Gibraltar) margins of western Tethys and also microplates within this extensional marine basin (Italy, Greece) (Fig. �). Continuous, well-ex-posed, stratigraphic sections have been identified and logged spanning the late Hettangian to early Pliensbachian time inter-val (Fig. 2a, b). The strata are limestones and dolomites with occasional shales that accumulated in peritidal, inner platform environments. All of the seven stratigraphic sections are char-acterised by high-frequency, peritidal cycles that are being de-scribed elsewhere (e.g. Bosence et al. 2006). The material for this study comes from the shallow subtidal facies of these cy-cles that commonly occur in shallowing-upward, but also deep-ening-upward, metre-scale cycles. Thicknesses, lithologies and foraminiferal records of the sections are given in figures � and 2 and they comprise: Buffedero Member, Gibraltar Limestone Formation, Gibraltar (Bosence et al. 2000; BouDagher-FaDel et al. 200�); Gavilàn Formation, Lomo Prieto, Betic Cordillera, Spain (ruiz-ortiz et al. 2004); Cuevas Labradas Formation, Iberian Range, Spain (coMas-rengiFo et al. �999); Jebel Rat Formation, Aghbalou n’Kerdous, southern High Atlas, Moroc-co (septFontaine �984); Oust Formation, Jebel Aziz, Dorsales Range, Tunisia (soussi 2002); Calcare Massiccio, Monte Bove south, Sibillini Mountains, central Italy (Bice & steWart �990; casaglia 2003/4); Pantokrator Formation Chalkis Quarry, Ev-via, Greece (scherreiKs 2000). The strata are limestones and dolomites with occasional shales that accumulated in peritidal, inner platform environments and the details of the facies and palaeoenvironments are to be published seperately. Figure 3 shows approximately where the studied sections accumulated in the Early Jurassic around �80 Ma. See Appendix for details of localities and ages of sequences.

The regional dataset therefore comprises a stratigraphically and sedimentologically well-constrained record from this time slice, within this critical period for the recovery and Mesozoic diversification of the benthic foraminifera. It traces the recov-ery of the benthic foraminifera from the Triassic-Jurassic crisis to the Pliensbachian and correlates the assemblages of North Africa with Spain, whilst emphasising the relative isolation within Tethys of the offshore Apuleian and Pelagonian plat-forms during this period (see Figure 3). From this regional study and comparison it is clear that five biozones can be recognised in this interval. In the past, these Early Jurassic, inner platform carbonates have proved difficult to date and this paper provides stage and sub-stage level dating of these Tethyan carbonate sections for the first time.

The study material consists of �92 random thin sections provided and logged by collaborators from different countries and institutions (see Appendix). Gibraltar material is deposited in the Natural History Museum, London.

Systematic palaeontology

Foraminifera

Superfamily P f e n d e r i n a c e a sMout & sugDen �962

Subfamily P s e u d o p f e n d e r i n i n a e septFontaine �988

Genus Pseudopfenderina höttinger �967

Pseudopfenderina cf. butterlini (Brun �962)Plate 7, fig. 7; Plate �0, fig. 6

R e m a r k s : A Pseudopfenderina with a high trochospiral and numerous chambers. The umbilical part of the chamber interior is filled with numerous pillars that are continuous from chamber to chamber.

D i s t r i b u t i o n :• Present work: Arghbalou n’Kerdous, southern High Atlas,

Morocco; Calcare Massiccio (Sinemurian – Pliensbachi-an), Monte Bove south, Sibillini Mountains, central Italy; Jebel Aziz, Dorsales Range, Tunisia; Almonacid da Cuba, Iberian Basin, Spain.

Genus Everticyclammina reDMonD �964

Everticyclammina praevirguliana Fugagnoli 2000Plate 3, fig. 6, Plate 4, fig. �, 5, Plate 5, fig. 3, Plate 9, figs 2-3

�984 Everticyclammina sp. nov. septFontaine p. 224, pl. �, figs 2-4.2000 Everticyclammina praevirguliana Fugagnoli p. �27, pls �-3

(cum. syn.).200� Everticyclammina praevirguliana Fugagnoli – BouDagher-

FaDel et al., p.6��, pl. 2, fig. �2.

D i m e n s i o n s : � to �.4 mm longest diameter.

R e m a r k s : An Everticyclammina with an alveolar test wall and single septal aperture. The initial coil is very short and missing in most random sections that show the elongate unise-rial part only.

Our material agrees closely with the uniserial tests il-lustrated and described by Fugagnoli (2000) from the Early Jurassic of northern Italy as Everticyclammina praevirguliana, the only Early Jurassic species known of this distinctive and otherwise largely Late Jurassic to Early Cretaceous genus. It is also very similar to the unnamed new species figured from Morocco by septFontaine (�984), whose specimens Fugagnoli (2000) includes in synonymy with her own species, together with specimens less confidently identified by septFontaine (�977) from France and by gušič (�977) from Yugoslavia. Ad-


Fig. 2a. Logged stratigraphic sections with foraminiferal distribution from Gibraltar, Iberia and Almonacid de la Cuba.


Fig. 2b. Logged stratigraphic sections with foraminiferal distribution from Morocco, Tunisia, Italy and Greece.


ditionally, we note that it is similar to a specimen figured by gonzález-Donoso et al. (�974, pl. 4, fig. 9) from the Lower Jurassic of southern Spain as an indeterminate lituolid with a complex internal structure. In the Eastern Mediterranean, nouJaiM clarK and BouDagher-FaDel (2004) recorded the first occurrence of Everticyclammina in the Middle to Upper Jurassic of Lebanon.

D i s t r i b u t i o n :• Literature: Lower Jurassic (Lower Lias) of Cabrera, one

of the Balearic Islands of Spain (coloM, �980); Lower Jurassic (Sinemurian-Pliensbachian) Morocco (septFon-taine �984); late Sinemurian-Pliensbachian age from the Rotzo Member, Calcari Grigi Formation, in the Trento Platform of the Venetian Prealps (Fugagnoli 2000); Mid-dle Jurassic of the French Prealps (septFontaine �977); Jurassic of Yugoslavia (gušič �977).

• Present work: Buffedero Member, Gibraltar Limestone Formation (Sinemurian), Gibraltar; Gavilàn Formation, Lomo Prieto (Sinemurian – early Pliensbachian), Betic Cordillera, Spain; Cuevas Labradas Formation (mid Sine-murian), Iberian Range, Spain; Jebel Rat Formation (Sine-murian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco; Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian); Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy; Pantokrator For-mation (Sinemurian – Pliensbachian), Chalkis Quarry, Ev-via, Greece (Figs �, 2).

Family M e s e n d o t h y r i d a e voloshinova �958

Subfamily O r b i t o p s e l l i n a e höttinger & caus �982

Genus Orbitopsella Munier-chalMas �902

Orbitopsella primaeva (henson �948)Plate �, figures �, 2, 4, Plate 2, fig. 4

1948 Coskinolinopsis primaevus henson, p.27, pl.�0, figs 4-5.1967 Orbitopsella primaeva (henson), höttinger, p.46, text-figs

23 (k-s), pl. 4, figs �7-�8.1984 Orbitopsella primaeva (henson), septFontaine p.2�4, pl.�,

fig.�2.1998 Orbitopsella primaeva (henson), Fugagnoli, p.�47, pl. �, figs

�-9, ol.2, figs �-�0.2004 Orbitopsella primaeva (henson), Fugagnoli, pl.�, fig.�.

D i m e n s i o n s : 4 to 5 mm greatest diameter in equatorial view.

R e m a r k s : An Orbitopsella with a discoidal test, first stage planispiral followed by a flaring, flabelliform stage with 35 to 40 annular chambers. The wall is with simple endoskeletal and exoskeletal pillars.

This species is smaller in size than Orbitopsella praecur-sor which reaches 8 to �0 mm, according to höttinger (�967). However, O. primaeva has a higher number of spiral chambers; 35 compared to �2 chambers.

Fig. 3. An Early Jurassic palaeogeographical map of Western Tethys modified after thierry et al. (2000) and DoMMergues et al. (2005) showing all studied locations, except Gibraltar as this is unknown.


D i s t r i b u t i o n :• Literature: Lower Jurassic of the Musandan limestone,

Oman (henson �948); Pliensbachian – Toarcian of Tur-key (septFontaine �984); upper Sinemurian – lower Pliensbachian of Morocco (Höttinger �967, septFontaine �98�, Bassoullet �998; Fugagnoli �998).

• Present work: Gavilàn Formation, Lomo Prieto (Sinemu-rian – early Pliensbachian), Betic Cordillera, Spain (Figs �, 2).

Orbitopsella praecursor (güMBel �872)Plate 3, fig.3

�872 Orbitolites praecursor güMBel p.256, pl.7, figs �-�0.�967 Orbitopsella praecursor (güMBel �872), höttinger, p.40,

text-fig. 20, pl.5, fogs �-�2.�994 Orbitopsella praecursor (güMBel �872), chiocchini et al.,

pl.2, figs �2-�3.�998 Orbitopsella praecursor (güMBel �872), Fugagnoli &

Broglio loriga, p.52, fig. 6.6-9 (see references).

D i m e n s i o n s : 4 to 5 mm greatest diameter in equatorial view.

R e m a r k s : This form is distinguished in having �2 spiral chambers in the microspheric forms and 8 to 9 in the mega-lospheric froms.

O. praecursor has a semi-circular test compared to the flar-ing test of O. primaeva and less delimited annular canals com-pared to O. dubari höttinger �967. Our specimens are mainly microspheric forms.

D i s t r i b u t i o n :• Literature: Uppermost Sinemurian - Plienbachian of Italy

(septFontaine �988, Fugagnoli & Broglio loriga �988).• Present work: Gavilàn Formation, Lomo Prieto (Sinemu-

rian – early Pliensbachian), Betic Cordillera, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco (Figs �, 2).

Genus Lituosepta cati �959

Lituosepta recoarensis cati �959Plate �, fig. 2, Plate 2, fig.3

1959 Lituosepta recoarensis cati, p. 3, pl. �, figs �-5, 7-��, ?�4.�962 Lituosepta recoarensis cati, sartoni & crescenti, p. 20, pl.

2, fig. 6.1967 Lituosepta recoarensis cati, höttinger, p.34-35, pl. 4, figs

�4-�6, pl.8, figs 27-29, txt-figs �5-�6.�969 Labyrinthina recoarensis (cati), gušič, p.63-64, pl.4, figs �-

2, pl. 5, fig. 5�998 Lituosepta recoarensis cati, Fugagnoli, p. �50, pl. 5, figs �-

8.2004 Lituosepta recoarensis cati, Fugagnoli, pl.�, fig. 8.

D i m e n s i o n s : 0.5 to �.4 mm longest diameter.

R e m a r k s : A Lituosepta with a flaring flabelliform test. The canaliculated wall distinguishes Lituosepta from Labyrinthina WeynschenK.

D i s t r i b u t i o n :• Literature: Lower Jurassic of Italy (cati �959); Sinemurian

of Morocco (höttinger �967), Turkey (septFontaine �98�, �984); upper Sinemurian (Fugagnoli �998, 2004).

• Present work: Gavilàn Formation, Lomo Prieto (Sinemu-rian – early Pliensbachian), Betic Cordillera, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco; Pantokrator Formation (Sinemurian – Pliensbachian), Chalkis Quarry, Evvia, Greece (Figs �, 2).

Lituosepta compressa (höttinger �967)Plate �, fig. 6, Plate 3, figs 2, 4

�967 Lituosepta compressa höttinger, p.36-38, pl. 4, figs �-�3.�988 Planisepta compressa (höttinger), septFontaine, p. 242.1994 Lituosepta compressa höttinger, chiocchini et al., pl.2,

fig. 8/1998 Lituosepta compressa höttinger, Fugagnoli, p.�52, pl. 6,

figs �-7.2004 Lituosepta compressa höttinger, Fugagnoli, pl.2, fig. 4.

D i m e n s i o n s : 0.4 to �.5 mm longest diameter.

D i a g n o s i s : This species is characterised by having a sim-ple proloculus followed by a planispiral stage, and a well de-veloped uncoiled part which becomes fan shaped, particularly in the microspheric forms. The test possesses multiple cribrate apertures. The apertural openings between the pillars penetrate the height of the chamber space. Peripheral partitions are seen in transverse sections.

R e m a r k s : septFontaine (�988) created a new genus Plani-septa for this species on the basis that it differs from Litu-osepta because of the lateral compression of the test.

D i s t r i b u t i o n :• Literature: Pliensbachian of Morocco, upper Pliensbachian

of Italy (Fugagnoli �998, 2004).• Present work: Gavilàn Formation, Lomo Prieto (Sinemu-

rian – early Pliensbachian), Betic Cordillera, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco (Figs �, 2).

Family H a u r a n i i n d a e septFontaine �988Subfamily A m i j i e l l i n a e septFontaine �988

Genus Amijiella loeBlich & tappan �985

Amijiella amiji (henson �948)Plate �, fig. 5, Plate 3, fig. 5, Plate 7, fig. �, Plate 8, fig. �

1948 Haurania amiji henson, p.�2, pl. �5, figs 5-�0.1998 Amijiella amiji (henson), Fugagnoli, p. �6�, pl. �2, figs �-9

(see list of references).2004 Amijiella amiji (henson), Fugagnoli, pl.2, fig. �.

D i m e n s i o n s : �.3 to �.5 mm longest diameter.

R e m a r k s : This species is characterised by having high chambers, septa very arched, initial coil very small. The unise-rial test has strong radial partitions, tending to become thick-ened (in transverse section) towards central zone, only rarely


bifurcate vertically to form few, scattered chamberlets. Earlier apertures are areal and multiple, but the last formed ones may be reduced in numbers or even become single.

The primary types of Amijella amiji from limestone dug out of a water-well at Wadi Amij, Iraq, were originally consid-ered (henson �948) to be of “Jurassic or Triassic age” as they were thought to have come from the Mulassa Limestone (a formation which is late Triassic in part). The primary types are now known to have come from the Jurassic of the Muhaiwir Formation (which has its type locality in Wadi Hauran, on the road to Qasr Amij.

D i s t r i b u t i o n :• Literature: Upper Sinemurian – Bathonian of Iraq (henson

�948), Turkey, Morocco (höttinger �967), Italy (Fugag-noli �998, 2004).

• Present work: Gavilàn Formation, Lomo Prieto (Sinemu-rian – early Pliensbachian), Betic Cordillera, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco (Figs �, 2).

Genus Haurania henson �948

Haurania deserta henson �948Plate 2, fig. 6, Plate 8, figs 2-5

�948 Haurania deserta henson p. �2, pl. �5, figa �, 3, 4�997 Haurania deserta henson, Banner et al., pl.�, fig. 9, �3, �4.�998 Haurania deserta henson, Fugagnoli, p.�7�. pl. �8, figs �-7.2004 Haurania deserta henson, Fugagnoli, pl. 2, figs 2, 3.

D i m e n s i o n s : � to �.5mm longest diameter.

R e m a r k s : This species is characterised by having delicate radial partitions, often bifurcating vertically to form a partial tier of peripheral chamberlets. The type specimens of H. deserta have been obtained from the Mulussa Limestone Formation, Iraq (upper Triassic in part), but are now believed to have prov-enance in the Muhaiwir Formation (Dunnington et al., �959), of ?early Bathonian age, which has its type locality in Wadi Hau-ran. Megalospheric forms have terminally high domed cham-bers and restricted areas for the terminal, cribrate aperture. Mi-crospheric forms are large with broader, flatter terminal faces.

D i s t r i b u t i o n :• Literature: Upper Sinemurian – Bathonian Iraq (henson

�948), Morocco (höttinger �967), Turkey (Bassoulet & poisson �975), Italy (Fugagnoli �998, 2004).


Superfamily B i o k o v i n a c e a gušiç �977Family B i o k o v i n i d a e gušiç �977

Genus Bosniella gušiç �977

Bosniella oenensis gušiç �977Plate 5, figs 4, 6, Plate 7, figs 3-4, Plate 8, fig. 6

�977 Bosniella oenensis gušiç, p.�3-�4, pl.��, figs �-2, pl.�2, figs �-4, pl. �3, figs �-3.

�998 Bosniella oenensis gušiç, Fugagnoli, p. �73, pl. �9, figs �-9, pl.20, figs �-2.

�998 Bosniella oenensis gušiç, Fugagnoli & Broglio lorigA, p.63, Fig. �0 �-5.

D i m e n s i o n s : 0.5 mm greatest diameter.

R e m a r k s : This species is characterised by having a well developed uncoiled later stage with thick, widely spaced and gently curved septa. The aperture is single in the coiled stage becoming cribrate in the uncoiled part.

The presence or absence of alveoles in the wall is still debatable. septFontaine (�988, p.242) put the genus Bosniella in synonymy with Mesoendothyra Dain, as “the presence or ab-sence of keriotheka which is not always visible due to diagen-esis, is not a reliable criterion for the distinction between the taxons Mesoendothyra and Bosniella”. The type species of Me-soendothyra was originally described as having alveoles within the wall structure. However, in our specimens the wall structure is solid and we consider the 2 taxons as being separate.

D i s t r i b u t i o n :• Literature: Middle Lias of the Dinarids (gušiç �977), up-

per Sinemurian-Pliensbachian of Italy (Fugagnoli �998, Fugagnoli & Broglio loriga �998).


Superfamily Ve r n e u i l i n a c e a cushMan �9��Family Ve r n e u i l i n i d a e cushMan �9��

Genus Siphovalvulina septFontaine �988

Siphovalvulina colomi BouDagher-FaDel, rose, Bosence & lorD 200�

Plate 6, fig. 6, Plate 9, figs 4-5, Plate �0, fig. �, Plate ��, figs 2,4

�980 Eggerellina sp; coloM, fig. 4. 28-33.200� Siphovalvulina colomi BouDagher-FaDel et al., p.605, pl.�,

figs �-4.


R e m a r k s : A Siphovalvulina with an elongate test of up to five whorls of chambers; three chambers in each whorl, the chambers oviform in shape with smoothly convex terminal faces and septa and high, convex apertural faces which sur-round a narrow umbilicus; weakly depressed sutures; and smooth test outline, the test becoming nearly parallel-sided with later growth.

Siphovalvulina colomi differs from Siphovalvulina vari-abilis septFontaine (�988) in possessing a much more com-pact test, only weakly depressed sutures, and smoothly convex septa which are not highly arched and oblique to the main axis as in S. variabilis. It differs from Siphovalvulina gibraltarensis BouDagher-FaDel et al. (200�) in possessing a less widely flaring test and more weakly depressed sutures.

D i s t r i b u t i o n :• Literature: Lower Jurassic (Lower Lias) of Cabrera, Balearic Is-

lands of Spain (coloM �980); Buffedero Member (Sinemurian),


Gibraltar Limestone Formation of Gibraltar (BouDagher-FaDel et al. 200�).

• Present work: Buffedero Member, Gibraltar Limestone Forma-tion (Sinemurian), Gibraltar; Gavilàn Formation, Lomo Prieto (Sinemurian – early Pliensbachian), Betic Cordillera, Spain; Oust Formation (early-mid Sinemurian), Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy; Pantokrator Forma-tion (Sinemurian – Pliensbachian), Chalkis Quarry, Evvia, Greece (Figs �,2).

Siphovalvulina gibraltarensis BouDagher-FaDel, rose, Bosence & lorD 200�

Plate 2, figs �,2, Plate 4, fig. 2A, B, Plate 6, figs 3-5, Plate 9,

fig.6, Plate ��, figs �, 5

�980 Eggerellina sp; coloM, fig. 4. 64, 66, 75, 76.200� Siphovalvulina gibraltarensis BouDagher-FaDel et al., p.605,

pl. �, figs 6-��.


R e m a r k s : A Siphovalvulina with widely flaring test of up to four rapidly enlarging whorls of chambers; three chambers in each whorl; chambers depressed in shape and with smooth-ly convex terminal faces but sharply truncated and flattened apertural faces, surrounding a broad umbilicus; and depressed sutures, producing a lobulate outline in both vertical and trans-verse sections.

Siphovalvulina gibraltarensis has a much more broadly flaring test (external angle of spire 900 or more), flatter aper-tural faces and broader umbilicus than either S. variabilis sep-tFontaine or S. colomi (BouDagher-FaDel et al. 200�).

D i s t r i b u t i o n :• Literature: Lower Jurassic (Lower Lias) of the Subbetic of Spain

(González-Donoso et al. �974) and of Cabrera, Balearic Islands (coloM �980); Buffedero Member (Sinemurian) of the Gibral-tar Limestone Formation of Gibraltar (BouDagher-FaDel et al. 200�).

• Present work: Buffedero Member, Gibraltar Limestone Formation (Sinemurian), Gibraltar; Gavilàn Formation, Lomo Prieto (Sine-murian – early Pliensbachian), Betic Cordillera, Spain; Cuevas Labradas Formation (mid Sinemurian), Iberian Range, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco; Oust Formation (early-mid Sinemurian), Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy; Pantokrator Formation (Sinemu-rian – Pliensbachian), Chalkis Quarry, Evvia, Greece (Figs �,2).

Genus Duotaxis Kristan �957

Duotaxis metula Kristan �957Plate 4, fig. 6

�957 Duotaxis metula Kristan, p. 294, pl. 27, figs 5a-d, 6.?�959 Tetrataxis conica ehrenBerg; Farinacci, p. �2, pl. 5, fig. 2.�964 Duotaxis metula Kristan; Kristan-tollMann, p. 46, pl. 7, fig.

�0.�980 Valvulina cf. metula (Kristan); coloM, fig. 4.�5-�6.�996 Duotaxis metula Kristan; Fugagnoli, p. 388, pl. �, figs �-5;

fig. 2a-h.

200� Duotaxis metula Kristan, BouDagher-FaDel et al., p. 606, pl.2, figs �-4.

D i m e n s i o n s : 0.3 to 0.4 mm greatest diameter

R e m a r k s : Originally regarded as a member of the Tetratax-inae, Duotaxis differs in having an agglutinated rather than a two-layered microgranular calcareous wall and was thus reassigned to the Family Verneuilinidae by loeBlich & tappan (�988). The genus had earlier (loeBlich & tappan �964) been considered a synonym of Valvulina, but differs in the absence of an early triangular stage and in lacking a truly valvular tooth (loeBlich & tappan �988).

D i s t r i b u t i o n :• Literature: Widespread in the Trias (?Ladinian-Rhaetian) to Lower

Jurassic (Pliensbachian), notably Rhaetian of Austria (Kristan �957; Kristan-tollMan �964), Lower Jurassic (Lower Lias) of Cabrera, Balearic Islands (coloM �980), and Pliensbachian of the Apennines, Italy (Farinacci �959), but also known from the south-ern Alps, southern Spain, and Morocco (fide Fugganoli �996); the Buffedero Member (Sinemurian) of the Gibraltar Limestone For-mation of Gibraltar (BouDagher-FaDel et al. 200�).

• Present work: Buffedero Member, Gibraltar Limestone Forma-tion (Sinemurian), Gibraltar; Gavilàn Formation, Lomo Prieto (Sinemurian – early Pliensbachian), Betic Cordillera, Spain; Cue-vas Labradas Formation (mid Sinemurian), Iberian Range, Spain; Oust Formation (early-mid Sinemurian), Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy; Pantokrator Formation (Sinemurian – Pliensbachian), Chalkis Quarry, Evvia, Greece (Figs �,2).

Superfamily S p i r o p l e c t a m m i n a c e a cushMan �927

Family Te x t u l a r i o p s i d a e loeBlich & tappan �982

Genus Textulariopsis Banner & pereira �98�

Textulariopsis sinemurensis new speciesPlate 5, fig. �, Plate 9, fig. �, Plate ��, fig.3

200� Textulariopsis sp. BouDagher-FaDel, rose et al., p.608, pl. 2, figs. 5-7.

M a t e r i a l : 36 specimens. Types figured here: H o l o t y p e P.66936, Pl.��, fig. 3, paratype pl. 9, fig. �.

D i m e n s i o n s : 0.3 to 0.4 mm greatest diameter Diagnosis. A Textulariopsis with an elongate test and with al-most parallel sides. Test may be slightly twisted.

R e m a r k s : On the basis of a single (type) species, Banner & pereira (�98�, p. 98) diagnosed their new genus Textu-lariopsis as ‘Wall agglutinated with calcareous cement, solid, imperforate, lacking canaliculi or pseudopores; proloculus succeeded by a rectilinear series of chambers, all biserially ar-ranged; aperture anteriomarginal, a simple basal, narrow slit. Differs from Spirorutilus hoFKer in its lack of a planispiral ini-tial stage, from Textularia DeFrance in its lack of canaliculate, pseudoporous walls, and from Pseudobolivina Wiesner by its

BouDagher-FaDel & Bosence: Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates...�0

low aperture and structurally insignificant chitinoid endoskel-eton’. Two new Cretaceous species were assigned to the genus by loeBlich & tappan (�982), plus three other Cretaceous spe-cies from the USA, and T. areoplecta from the Lower Jurassic (Pliensbachian-Toarcian) of northern Alaska which had previ-ously been assigned to Textularia. Our specimens conform to the generic diagnosis, but cannot be matched closely either with T. areoplecta or any of the Cretaceous species.

Textulariidae have been reported as locally common in the Mediterranean Lias, e.g. in association with siphovalvulines in the Lower Lias of southern Spain (gonzález-Donoso et al. �974) and northern Italy (Fugagnoli 2000) but without specific identification. It seems probable that some of this material may prove to be congeneric and possibly conspecific with our specimens. Thus although gonzález-Donoso et al. (�974, pl. 2, figs 5, 8) tentatively identified material from the Subbetic of southern Spain as Palaeotextularia?, their figured specimens have the solid wall structure of Textulariopsis rather than the two-layered wall structure characteristic of Palaeotex-tularia, a typically Carboniferous genus. However, since the wall structure is largely unknown for other potentially com-parable Jurassic species currently assigned to Textularia or similar genera, it is likely that some earlier records belong to our new species.

The Mediterranean specimens constitute the first record of Textulariopsis as such from the Mediterranean region, and confirm that this seemingly largely Cretaceous genus had originated by the Early Jurassic.

D i s t r i b u t i o n :• Literature: Buffedero Member (Sinemurian) of the Gibraltar

Limestone Formation of Gibraltar BouDagher-FaDel et al. (200�); Lebanon (nouJaiM clarK & BouDagher-FaDel 2003).

• Present work: Buffedero Member, Gibraltar Limestone Forma-tion (Sinemurian), Gibraltar; Gavilàn Formation, Lomo Prieto (Sinemurian – early Pliensbachian), Betic Cordillera, Spain; Cue-vas Labradas Formation (mid Sinemurian), Iberian Range, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco; Oust Formation (early-mid Sinemurian), Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy; Pantokrator Formation (Sinemu-rian – Pliensbachian), Chalkis Quarry, Evvia, Greece (Figs �, 2).

Superfamily Te x t u l a r i a c e a ehrenBerg �838Family C h r y s a l i d i n i d a e neagu �968

Genus Riyadhella reDMonD �965

Riyadhella praeregularis BouDagher-FaDel, rose, Bosence & lorD 200�

Plate 4, fig. 4, Plate �0, fig. 2, Plate ��, fig. 6

200� Riyadhella praeregularis BouDagher-FaDel et al., p. 6�0, pl.2, figs 8-��.

D i m e n s i o n s : 0.3 to 0.4 mm greatest diameter

R e m a r k s : A small species of Riyadhella characterised by thin and very strongly convexly curved septa, and a canalicu-late test which is quadriserial in neanic and ephibic as well as nepionic stages.

Riyadhella praeregularis appears to be an Early Jurassic (Sinemurian-Lower Pliensbachian) ancestor of the Mid Jurassic (Bathonian-Callovian) Riyadhella regularis reDMonD. Externally the two species are virtually identical. The cham-bers of R. praeregularis are very similar in shape to those of R. regularis as illustrated in sections by Banner et al. (�99�, p. �30, fig. 57d and p. �32, fig. 63e), but the septa are much thinner and more convexly curved in R. praeregularis. In R. regularis the early stage of growth is quadriserial but becomes triserial in the adult, whereas R. praeregularis is quadriserial throughout.

R. arabica reDMonD is a poorly known species, but larger than R. praeregularis. R. elongata reDMonD is easily distin-guished by its thin, parallel-sided test and its ontogenetically late development of triseriality, R. inflata reDMonD by its large, thick test and adult triseriality. Banner et al. (�99�, fig. �05) il-lustrated the descent of R. regularis from Bajocian-Bathonian R. elongata via early Bathonian R. arabica; our Sinemurian R. praeregularis pre-dates all these species, so is the most likely common ancestor yet known.

In having more highly convex septa and terminal faces, R. praeregularis differs from species of Redmondoides Banner et al. (�99�), although this genus has initially solid walls which may become canaliculate in the last-formed chambers, and the test although quinqueserial or quadriserial initially becomes quadriserial in the adult.

Although Banner et al. (�99�) stated that no canaliculation or canaliculi are known for the type or other species of Riyad-hella, they described and discussed the common development of canaliculi in other chrysalidinid genera. Recognition herein of canaliculi in R. praeregularis is therefore consistent both with the amended diagnosis of Banner et al. (�99�) for the Family Chrysalidinidae and their ascription of Riyadhella to it.

D i s t r i b u t i o n :• Literature: Hitherto the earliest record of Riyadhella was from the

Mid Jurassic (late Bajocian) Dhruma Formation of Saudi Arabia (reDMonD �965; Banner et al., �99�). Description of Riyadhella from the Buffedero Member (Sinemurian) of the Gibraltar Lime-stone Formation by BouDagher-FaDel et al. (200�) therefore extended the known stratigraphic range of the genus down into the Lower Jurassic, and its geographical range from the Middle East (Saudi Arabia and the United Arab Emirates), Switzerland, and possibly Siberia (fide Banner et al. �99�) to include the west-ern Tethys. In the Middle East it extends into the Upper Jurassic (Kimmeridgian).

• Present work: Buffedero Member, Gibraltar Limestone Formation (Sinemurian), Gibraltar; Gavilàn Formation, Lomo Prieto (Sine-murian – early Pliensbachian), Betic Cordillera, Spain; Cuevas Labradas Formation (mid Sinemurian), Iberian Range, Spain; Jebel Rat Formation (Sinemurian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco; Oust Formation (early-mid Sinemurian), Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy; Pantokrator Formation (Sinemu-rian – Pliensbachian), Chalkis Quarry, Evvia, Greece (Figs �,2).

P r o b a b l e c y a n o b a c t e r i u m

Genus Cayeuxia Frollo �938

Ty p e s p e c i e s : Cayeuxia piae Frollo �938, from the Upper Jurassic of the eastern Carpathian mountains, eastern Europe.

BouDagher-FaDel & Bosence: Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates... ��

Cayeuxia ?piae Frollo �938Plate 6, fig.�, Plate 7, fig. 5, Plate �0, fig. 3

�938 Cayeuxia piae Frollo, pp. 269-27�, pl. �8.�99� Cayeuxia piae Frollo; oKla, p. 93, pl. 3, figs �-3 (cum syn.).

R e m a r k s : Tufts of thin (approximately 0.� mm) closely packed filaments, without cross-partitions, are assigned to Cayeuxia, a taxon currently interpreted as a calcified cy-anobacterial sheath (riDing �99�) comparable and arguably synonymous with the present-day Rivularia (cf. Dragastan �985, �988). The distinction and generic assignment of over a dozen species previously assigned to Cayeuxia were re-viewed by Dragastan (�985), but since species diagnostic characters have proved controversial, we provisionally adopt the species name by which such microbes are widely known in the Mediterranean Lias (fide sartorio & venturini �988). The specimens of Cayeuxia differ significantly from those of Stromatomorpha liasica described from the north face of Gi-braltar by le Maître (in DuBar & le Maître �935, pl. 8, figs �-5). These ‘spongiomorphids’ (= stromatoporoid sponges, fide oWen & rose �997) have a more complex, larger-scale internal structure with clear cross-partitions.

D i s t r i b u t i o n :• Literature: Cayeuxia and probable synonyms have been widely re-

ported in circum-Mediterranean countries, from at least the Mid-dle Trias to Lower Cretaceous (Dragastan �985); C. piae ‘very common in the Lias, is present throughout the Jurassic and in the Early Cretaceous’ (sartorio & venturini �988, p. 68). Similar forms have been illustrated from the Lower Jurassic Pantokrator Limestones of Corfu, Greece (Flügel �983, p. 268, pl. 48), from the Lower Jurassic (Hettangian-Sinemurian) Calcare Massiccio of the central Apennines, Italy (Barattolo & Bigozzi �996, pl. 43, fig. �; pl. 44, fig. 2), from the Lower Jurassic of the Balearic Is-lands (coloM �980, fig. 2. 23), and also from the Middle Jurassic (Callovian) Tuwaiq Mountain Limestone of Saudi Arabia (oKla �99�).

• Present study: Buffedero Member, Gibraltar Limestone Forma-tion (Sinemurian), Gibraltar; Cuevas Labradas Formation (mid Sinemurian), Iberian Range, Spain; Jebel Rat Formation (Sine-murian – Pliensbachian), Arghbalou n’Kerdous, southern High Atlas, Morocco; Oust Formation (early-mid Sinemurian), Jebel Aziz, Dorsales Range, Tunisia; Calcare Massiccio (Sinemurian – Pliensbachian), Monte Bove south, Sibillini Mountains, central Italy.

A l g a ?

Genus Thaumatoporella pia �927

Ty p e s p e c i e s : Gyroporella parvovesiculifera raineri �922, from the Upper Cretaceous of Italy

Thaumatoporella ?parvovesiculifera (raineri �922)Plate 2, fig. 5, Plate 4, fig. 3A, Plate 9, fig.7, Plate �0, fig.4,

Pl.��, fig. �0.

�922 Gyroporella parvovesiculifera raineri, p. 83, pl. 3, figs �7-�8.�962 Thaumatoporella parvovesiculifera (raineri); sartoni &

crescenti, p. 270, pl. �6; pl. 46, figs 2, 3, 5. �99� Thaumatoporella parvovesiculifera (raineri); oKla, p. 93, pl.

3, figs 4-6 (cum syn.).

R e m a r k s : Thin micrite-walled polygonal and ellipsoidal to spheroidal microfossils with indistinct wall perforations are here assigned to Thaumatoporella.

These forms commonly occur as concentrically arranged layers whereby smaller bodies are enclosed in larger bodies. Our material compares closely with the thaumatoporellids illustrated by Flügel (�983, pl. 49) from the Lias of Corfu, Greece, and by Barattolo & Bigozzi (�996, pl. 44) from the Lias of the central Apennines, Italy. Flügel inferred that thaumatoporellids should be interpreted as organisms with animal affinities, but other authors generally have followed raineri (�922) and pia (�927) in assigning them to the algae, most commonly to the green algae, and De castro (�990) has reviewed arguments for elevating them to ordinal status within this category. Our material compares closely with the type spe-cies as commonly illustrated from the Mediterranean Lias (e.g. by sartorio & venturini, �988, p. 68).

D i s t r i b u t i o n :• Literature: Thaumatoporella in general are widespread in the

Mesozoic (from the Upper Trias onwards) in carbonate platform facies (Barattolo �99�), and extend to the Paleocene (De cas-tro �990). The genus Thaumatoporella is especially common in Liassic platform carbonates of the southern Tethys (sartorio & venturini �988), notably the Pantokrator Limestones of Corfu, western Greece (Flügel �983) and the Calcare Massiccio of Italy (Barattolo & Bigozzi �996). Thaumatoporella parvovesiculifera has often been recorded in association with Palaeodasycladus mediterraneus (see below), e.g. in the western Venetian prealps (Bosellini & Broglio loriga �97�), and in the central Apennines – where it has been used as a zonal index within the Lower Lias (chiocchini et al. �994).

• Present study: It occurs commonly in all the sections analysed (see Fig. 2) and individual occurrences are not further recorded here.

D a s y c l a d a c e a n a l g a

Genus Palaeodasycladus pia �927

Ty p e s p e c i e s : Palaeocladus mediterraneus PIA, �920, from the Lower Jurassic (Middle Lias) of southern Italy.

Palaeodasycladus ?mediterraneus (pia �920)Plate 5, fig. 5, Plate 6, fig. 2, Plate 7, fig.6, Plate ��, figs 8,9

�920 Palaeocladus mediterraneus pia, p. ��8, text-fig. 22, pl. 6, figs �-5.

�927 Palaeodasycladus mediterraneus (pia); pia, pp. 78-79, fig. 62.

�962 Palaeodasycladus mediterraneus (pia); sartoni & crescenti, p. 269, pl. �3, fig. 2; pl. 46, fig. �.

�978 Palaeodasycladus mediterraneus (pia); Bassoullet et al., p. �92, pl. 22, figs 8-9; pl. 23, figs �-2 (cum syn.).

�983 Palaeodasycladus mediterraneus (pia); Flügel, p. 27�, pl. 46, figs 5-9; pl. 47, figs �-6.

R e m a r k s : Palaeodasycladus mediterraneus is a well-known, dasycladacean green alga. Bassoullet et al. (�978) gave a detailed illustrated account of the genus and its three assigned species, supported by an extensive bibliography. Our material is only tentatively identified as P. mediterraneus because, although it corresponds well with specimens widely


figured under this name by earlier authors (notably that illus-trated as typical of this species from the Mediterranean Lias by sartorio & venturini (�988, p. 69), it is insufficient to reveal the branching angles generally deemed to be species diagnos-tic (e.g. by Bassoullet et al. �978). Moreover, Barattolo & Bigozzi (�996) noted that the taxon has been the object of a debate concerning its specific status and generic characteristics (cf. DeloFFre & laaDila �990).

D i s t r i b u t i o n :• Literature: P. mediterraneus is common in the Lower Jurassic of

western Tethys, e.g. Morocco (fide DeloFFre & laaDila �990; but not Barattolo & Bigozzi �996), Italy (Flügel �99�; Barattolo & Bigozzi �996), Greece (Flügel �983; zaMBetaKis-leKKas �995), and the Subbetic of southern Spain (anDreo et al., �99�; rey �997), typically in lagoonal and shelf margin facies (Barattolo & Bigozzi �996) of Hettangian-Pliensbachian age (Bassoullet et al. �978). Bassoullet et al. (�978) accepted older records from Spain (southeast and Balearic regions), Morocco, Algeria, Tunisia, Italy, the former Yugoslavia, Albania, Greece, Turkey, Oman, and southern Iran, noting that its wide distribution was favoured by the widespread occurrence of shallow-water carbonate platforms in the Lower and Middle Lias. A zone of P. mediterraneus was

distinguished by sartoni & crescenti (�962) within the Lias of southern Italy, and by authors such as chiocchini et al. (�994) within that of central Italy. The genus as a whole is confined to the Lias (Barattolo �99�).

• Present study: P. ?mediterraneus occurs locally throughout all the sections illustrated in Fig. 2 but individual occurrences are not noted for reasons of space.

Biostratigraphy

Despite their common occurrence in the sections described in this work Lower Jurassic (Hettangian-Sinemurian) foramin-ifera from western Tethys are poorly known in the literature. Although septFontaine (�98�) proposed a standard Jurassic biozonation for the Mesogean realm on the basis of 56 forami-niferal generic ranges which took account of work by earlier authors, his scheme began with an Orbitopsella range zone ap-proximately equivalent to the Pliensbachian. None of his gen-era were confidently recorded from the underlying Sinemurian. Later, septFontaine (�984) recorded ‘Siphovalvulina’ from the Sinemurian-Pliensbachian (Domerian) of Morocco and dis-

Fig. 4. Distribution chart of the 5 biozones in the Western Mediterranean.

BouDagher-FaDel & Bosence: Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates... �3

tinguished a biozone of ‘Siphovalvulina’ and Mesoendothyra characterized by a fauna of small, primitive lituolids: the two index genera plus Everticyclammina sp. nov., Glomospira sp. and Earlandia sp.. These taxa occurred in all six biozones he distinguished in the Moroccan Lower and Middle Lias, but the upper boundary of this ‘interval zone’ was defined by the first appearance of Lituosepta recoarensis CATI, the index fossil for the overlying zone. Subsequently, Bassoullet (in cariou & hantzpergue �997) listed Involutina liassica as the only form that is present in the Hettangian on the shallow platforms, while Everticylammina spp. appears in the highest part of the lower Sinemurian. He also stated that Lituosepta recoarensis preceeded Orbitopsella primaeva in the upper Sinemurian. More recently, Fugagnoli (2004) reported that the Hettangian – lower Sinemurian interval belongs to ?Lituosepta recoarensis Zone while the upper part of the Sinemurian and Lower Pliensbachian to the Orbitopsella zone and finally the Upper Pliensbachian to the Lituosepta compressa zone.

In this study, samples from seven continuously ex-posed and complete sections from the Mediterranean region (Figs �, 2) can be used as a biostratigraphical and palaeo-geographic framework to establish new benthic foraminiferal biozones. In all seven areas the foraminifera are mostly tex-tulariids, with simple wall structures in the Hettangian and lower Sinemurian, becoming more complex internally with pillars and alveoles in the walls towards the upper Sinemurian and Pliensbachian. Most recorded species are long – ranging but their first appearance is used here to define and trace the biozones (Fig. 4):

– The oldest proposed biozone is the Siphovalvulina gibral-tarensis biozone of Hettangian age and this is present in Gibraltar (samples K3-L7), Almonacid de la Cuba, north-east Spain (samples 39, 5-�5), Tunisia (samples �6-33) and Greece, Evvia (samples 5-�4), Chalkis section (samples 49-56). This biozone belongs to a carbonate platform en-vironment characterized by algal laminites, oncolites, and other features of peritidal/lagoonal facies with a noticeable absence of more advanced textularids. Foraminifera are very rare and include simple Textularia spp. and Siphoval-vulina spp. (see Figs 2a, b);

– The Siphovalvulina colomi biozone corresponds to the lower to mid Sinemurian and includes S. gibraltarensis, Duotaxis metula, Riyadhella praeregularis in Gibraltar (samples L6, G�-G5), Almonacid de la Cuba, northeast Spain (samples �9 - 2�), Morocco (samples 27- 28), Tuni-sia (samples 48-84), and Greece, northeast Evvia (samples 59-87), Chalkis section (samples 48-�0�). Foraminifera are still rare in this interval but diversity has slightly increased from the Hettangian. I. liassica having made its first ap-pearance in the previous biozone is joined by Pseudop-fenderina cf. butterlini (Pl. 7, fig. 2; Pl. �0, fig. 6) which is common in this zone (see Figs 2a, b);

– Everticyclammina praevirguliana biozone corresponds to the mid Sinemurian and is present in Gibraltar (samples G8-G27), Almonacid de la Cuba, northeast Spain (samples 27-44), Lomo Prieto, southern Spain (samples ��-��4) Morocco (sample 29), Tunisia (sample 85), Monte Bove, Italy (samples �3-66) and Greece, northeast Evvia (sam-ples 90-�0�), Chalkis section (sample �02). It coincides with the first appearance of E. praevirguliana and includes

Siphovalvulina colomi, S. gibraltarensis, Textulariopsis sinemurensis, Riyadhella praeregularis, and Duotaxis met-ula. Our foraminifera of this biozone are conspecific with those illustrated by coloM (l980) from the Balearic island of Cabrera. His specimens were obtained from the Lower Lias (therefore potentially within the range Hettangian to lower Pliensbachian), from micritic limestones similar to those associated with the grainstones and packstones sam-pled on Gibraltar, and with similar associated algae/cyano-bacteria (Palaeodasycladus mediterraneus, Thaumatopore-lla and Cayeuxia). Elements of this fauna (corresponding to S. gibraltarensis, T. sinemurensis, and Everticyclam-mina praevirguliana as identified herein) have also been illustrated although not confidently identified from the Lower Lias in the Subbetic region of southern Spain (e.g. gonzález-Donoso et al. �974), and E. praevirguliana (as-sociated with Duotaxis metula, Siphovalvulina, and sup-posed textulariids) has been recorded more precisely from the Lower-Middle Lias (late Sinemurian-Pliensbachian) of northern Italy (Fugagnoli 2000) (see Figs 2a, b);

– The Lituosepta recoarensis and Orbitopsella spp. biozone is assigned to the upper Sinemurian. It coincides with the first appearance of L. recoarensis and includes Si-phovalvulina sp., Haurania deserta, Orbitopsella praecur-sor, Amijiella amiji, Pseudopfenderina sp., and Bosniella oenensis and is present in Lomo Prieto, southern Spain (sample �3�), the Arghbalou n’Kerdous section, Morocco (sample 49), Greece, northeast Evvia (samples �02-�06b), and Chalkis section (samples �02-�06);

– The Lituosepta compressa biozone corresponds to the low-er Pliensbachian and coincides with the first appearance of L. compressa. It includes Pseudocyclammina sp., Haura-nia deserta, Amijiella amiji, Riyadhella sp., Siphovalvulina sp., Siphovalvulina colomi, Siphovalvulina gibraltarensis, Textulariopsis sinemurensis, Duotaxis metula. Everticycla-mmina praevirguliana, Pseudocyclammina sp., Orbitopsel-la sp., Haurania sp., Lituosepta recoarensis, Orbitopsella “circumvulvata”, Textularia sp., Siphovalvulina sp., Or-bitopsella praecursor, Bosniella oenensis, Amijiella amiji, Haurania deserta, Pseudofenderina sp., Buccicrenata sp. and small miliolids. This zone is found in Lomo Prieto in the Betics (samples �35-�59), Morocco (samples 52-56) and Greece, northeast Evvia (samples �07b-��9) (see Figs 2, 3).

The algal/microbial taxa are also consistent with a Hettang-ian – Pliensbachian age range for the foraminiferal biozones. Palaeodasycladus mediterraneus is typically Hettangian to Pliensbachian in range (sartoni & crescenti �962; Bassoul-let et al. �978), as is the genus as a whole. Thaumatoporella parvovesiculifera and Cayeuxia piae have longer stratigraph-ic ranges but commonly co-exist with P. mediterraneus in the Hettangian-Sinemurian (cf. sartorio & venturini �988; Barattolo & Bigozzi �996, BouDagher-FaDel et al. 200�). P. mediterraneus, T. parvovesiculifera, C. piae and Siphovalvuli-na sp. range together through Hettangian-Sinemurian carbon-ates of the Calcare Massiccio of central Italy (chiocchini et al. �994, fig. 22), indicating a close similarity in age between these dolomitic limestones and those of the Gibraltar Lime-stone. A correspondingly close similarity in facies between the two sequences has been noted by Bosence et al. (2000).


Diversity and biotic recovery

The climatic crisis at the Triassic-Jurassic boundary contrib-uted to the mass extinction in the marine realm at the end of the Triassic (hallaM �996) and affected all groups of larger benthic foraminifera, notably the small survivors of the fu-sulinids, the involutinids and the miliolids. In the Hettangian, occurrences of small opportunistic genera such as Duotaxis dominate sparse assemblages of new small textulariid forms, such as Siphovalvulina and are here represented by the S. gi-braltarensis biozone.

In the early Sinemurian most assemblages were poor, dominated by small textulariids Siphovalvulina gibraltarensis, S. colomi, Textularia sp., Duotaxis metula, Riyadella praeregu-laris, Textulariopsis sinemurensis and small miliolids. These forms co-existed with Thaumatoporella ?parvovesiculifera and are represented here by the S. colomi biozone which ex-tends from Spain to North Evvia in Greece. The diversity of foraminifera is generally poor in this biozone and this reflects the more restricted facies (including early diagenetic dolo-mites) in the lower parts of the sections in Gibraltar, Lomo Prieto, Almonacid da Cuba and Morocco compared to the mid to upper parts of these sections.

From middle to upper Sinemurian, forms with internal pil-lars, such as Pseudopfenderina, and fine alveoles in the walls, such as Everticyclammina species, began to evolve in the Western Tethys. But it is not untill the upper Sinemurian that a full foraminiferal recovery became obvious in Western Tethys. This recovery is mirrored by an increase in new genera, as new forms evolved to fill the niches that had been vacated by the end Triassic extinction event. Orbitopsella praecursor, Ami-jiella amiji, Haurania deserta, Pseudopfenderina sp., Everti-cyclammina sp., Lituosepta recoarensis, Pseudopfenderina cf. butterlini, Bosniella oenensis appear in Morocco, with a similar but less diverse assemblage appearing in Spain at the same time. Such assemblages consist of benthic foraminifera with large test wall surface areas such as Orbitopsella. This genus has many small chamberlets which may have been used for farming endosymbionts (höttinger �982). Such larger fo-raminifera are highly adapted to mesotrophic and oligotrophic (nutrient–deficient) conditions (Fugagnoli 2004). Orbitopsella together with Lituosepta, Pseudopfenderina, Everticyclam-mina and Haurania indicate an inner platform environment in Morocco and Spain, they are usually encountered in shallow-water carbonate facies all along the southern Tethyan margin (septFontaine �984). Such an assemblage contrasts with the Hettangian and early Sinemurian small foraminifera, such as Textularia and Siphovalvulina, which are widespread on these platforms and have been considered as more tolerant of chang-ing conditions (septFontaine �984; Fugagnoli 2004) as might be expected to occur in these high frequency cycles.

The Sinemurian is also characterised by the occurrence of oncoids. These concentrically structured micritic grains have both radially and tangentially oriented microbial filament moulds (Bosence et al. 2000, fig. 5E). This co-occurrence rein-forces previous interpretations (e.g. Flügel �979; hug �998) that in the early Mesozoic oncoids occurred in relatively open-marine waters in inner platform settings. Comparable facies have been illustrated and described from the Lower Jurassic of Italy; in the north (Bosellini & Broglio loriga �97�; castel-larin �972), south (sartoni & crescenti �962) and central (e.g. chiocchini et al. �994) regions.

The foraminiferal recovery continued through to the Pliensbachian with no major extinction at the boundary. However, more evolved taxa became established during the Pliensbachian by filling the empty niches of the Sinemu-rian with new forms of lituolids. These went on to play an important role in the evolutionary development of this group throughout the Jurassic and Cretaceous. The early Pliensbachi-an in Western Tethys is represented by the L. compressa biozone. This biozone is seen in northern Spain, Morocco and in Greece. Dasyclad algae also became abundant and flour-ished in these shallow-marine carbonate environments, where foraminifera became large and developed blind chamberlets (alveoles) in the test walls to house symbiotic algae. Such features developed into forms with larger and more consistent alveoles in the Pliensbachian to Late Jurassic and Cretaceous, becoming very important components of Mesozoic rocks. In southern Spain foraminifera were diverse and closely compa-rable with those of Morocco in northern Africa. It is not sur-prising that these two localities had similar assemblages and facies during the Pliensbachian; they were contiguous and lo-cated within the same enclave of western Tethys. On the other hand, some algae and benthic foraminifera are rare in Greece and the Apennines. This could reflect the relative isolation of the offshore Apuleian and Pelagonian microplates during the Pliensbachian. These isolated microplates were surrounded by deeper water areas, and the foraminifera lived in the shallow-water carbonate platforms that were sampled in our logged sections. Clearly these areas had lower diversity faunas than the shallow carbonate shelves of the northern and southern Tethyan margins

In summary, benthic foraminiferal biozones appear to be the only form of biostratigraphic correlation that can be used for these the low-latitude carbonate platforms which dominated the western margins and microplates of Tethys during the Early Jurassic. Five new biozones are erected from the consist-ent appearance of taxa of benthic foraminifera, some of which are new species, others new records for the Early Jurassic, or for this region. The stratigraphical occurrence of the taxa is based on bed-by-bed logging of continuously exposed sec-tions from the northern and southern Tethys margins and from the Apuleian and Pelagonian microplates from within Tethys. The foraminiferal occurrences permit correlation of sections from the southern and northern shelf-margin of Tethys, whilst reflecting the relative isolation of the Apuleian and Pelagonian microplates within Tethys.

The foraminifera show a progressive diversification of forms through the late Hettangian to early Pliensbachian inter-val that is interpreted to reflect the evolutionary recovery and diversification of these biotas following the Triassic-Jurassic extinction event.

Acknowledgements

Funding for this project was provided by grants to Bosence from the Royal Society and from the Central Research Fund of the University of London; this support is gratefully acknowl-edged. The sections that provided the material for this study were identified and logged with collaborators from each of the following countries and institutions and this collaboration and supply of material for study has been essential to the suc-


cess of this project. We therefore thank J. rey (Depto. Geolo-gia, E.U.P Linares, Universidad de Jaén, Spain), L.M. nieto (Depto Geologia, Facultad de Ciencias Experimentales, Uni-versidad de Jaén, Spain), M. aurell and B. BaDenas (Depto. Ciencias de la terra, Universidad Zaragoza, Zaragoza, Spain), S. cirilli and F. casaglia (Dipartimento Scienze della Terra, Piazza Universita, Perugia, Italy), R. scherreiKs (Geologische Staatssammlung, Luisenstr. 37, Munich, Germany, for work in Greece), M. soussi and A. Bel Kahla (Departément de Géolo-gie, Université de Tunis, Tunis, Tunisia), M. MehDie (Departé-

ment de Géologie, Université Ibn Tofail, BP �33, Kenitra, Mo-rocco), and E. procter (Geology Department, Royal Holloway University of London, Egham, UK) for collaborative work in Tunisia. Research in Greece was carried out under permit from IGME, Athens and access to Chalkis Quarry with kind permission and assistance from the Halkis Cement Company of the Heracles Group, Chalkis. We are especially thankful for the careful editing and reviewing by Prof. Alan lorD (For-schungsinstitut Senckenberg), Prof. Roberto rettori (Piazza Università, Italy) and Prof. Malcolm hart (Plymouth, UK).

Appendix

Locations of logged sections. These are given approximately in Figs �, 3 and details only are given here.

Gibraltar. All details of sections and samples are given in BouDagher-FaDel et al. (200�). Only the Main Ridge section is illustrated here (Fig. 2) for comparison.

Lomo Prieto, Murcia Province, southern Spain. The location of this section in the Lower Member (M�) of the Gavilán For-mation is given in ruiz-ortiz et al. (2004, fig. 3). Brachiopods at the top of this section have been dated at �89.2 ma (early Pliensbachian) using 87/86 Strontium isotopes (ruiz-ortiz et al., 2004).

Almonacid de la Cuba, northeast Spain. This section is situ-ated in the baranco to the north of the village of Almonacid de la Cuba on the southern limb of a northwest-southeast trending anticline. It corresponds to the lower part of the section (0-�4m approximately) described by coMas-rengiFo et al. (�999) and also the exposed, but previously unlogged, section beneath.

Aghbalou n’ Kerdous, southern High Atlas, Morocco. This sec-tion is continuously exposed in the wadi north of the village of Aghbalou n’Kerdous. The log was measured on the northwest-ern side of the wadi along the irregation channel. The section is mainly within the Jebel Rat Formation with the exception of the top few metres that we interpret as the Aghbalou Forma-tion as described in the Carte Géologique du Maroc.

Jebel Aziz, Dorsales Range, Tunisia. This previously unlogged section in the Oust Formation (soussi, 2002) is on an isolated hill to the west of the town of Bir M’cherga (Fig. �). The meas-ured section lies between two quarries, an active one to the north and a disused one to the south and starts above the fields

at the base of the hill and continues to the crest of the hill where the overlying Zaghouan Formation is exposed. These overlying hemipelegic mudstones have been dated at this lo-cality by Fauré & peyBernes (�986) by ammonites indicating an early Pliensbachian (Tropidoceras zone) age.

Monte Bove South, Sibillini Mts. Central Apennines, Italy. This section is located to the on Monte Bove South to the west of the top of the ski lift that starts at the Hotel Felicitas, in Fontignano. The section, which is all within the Calcare Mas-siccio, was logged from the track leading west from the ski lift and continues to the summit of Monte Bove south and details are given in casaglia (2003/4), No samples have been stud-ied from above �00 m on the measured section illustrated in Fig. 2.

Chalkis Quarry, Chalkis, Greece. The very large cement quarry to the north of the road on the Athens side of the new Chalkis bridge provides a continuous, but sometimes faulted section through Lower Jurassic carbonates of the Pantokrator Formation. The measured section (Fig. 2) comes from a rela-tively unfaulted section that in 2002 was in the centre of the large quarried face to the west of a large faulted, brecciated and dolomitised unit. The continuous �20 m section measured lies between two faults.

In addition to the Chalkis section we illustrate (Plate 5) mate-rial from Mt Xiron Orus in northeast Evvia (Fig.�, scherreiKs, 2000). The samples come from the Pantokrator Limestone as follows: 4.��.� a large limestone oucrop SE of Mount Psili Rachi, between the village of Kerasia and (mount) Xiron Oros. �2.�0.6� and �2.�0.64 Eastern flank of Xiron Oros along for-est road at about 700 m elevation (pers com. r. scherreiKs, 2006).

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Plate �

All thin-section photomicrographs from Gavillán Formation, Lomo Prieto, Betic Cordillera, Southern Spain.

Figs �, 2, 4. Orbitopsella primaeva (henson �948), microspheric forms, sample RA-0�-�86, x22.Fig. 3. Lituosepta recoarensis cati, microspheric forms, sample RA-0�-�22, x54.Fig. 5. Amijiella amiji (henson �948), sample RA-0�-�80, x8�.Fig. 6. Lituosepta compressa (höttinger �967), sample RA-0�-�79, x53.

Manuscript received: 2006-�2-�5; accepted: 2007-05-�5

Senckenbergiana lethaea, 87 (�); 2007 Plate �

M. K. BouDagher-FaDel & D. W. J. Bosence: Early Jurassic benthic foraminiferal diversification and biozones in shallow-marine carbonates of western Tethys.


Plate 2


Figs �, 2. Siphovalvulina gibraltarensis BouDagher-FaDel et al. 200�, �. Vertical section sample RA-0�-�39, x �33. 2. Oblique transverse sections, sample RA-0�-�87, x83.

Fig. 3. A. Siphovalvulina sp., B, Lituosepta recoarensis CATI, megalospheric forms, RA-0�-�86, x66.

Fig. 4. Orbitopsella primaeva (henson �948), sample RA-0�-�59, megalospheric form, x 33.Fig. 5. Thaumatoporella ?parvovesiculifera (raineri �922), sample RA-0�-��, x33.Fig. 6. Haurania deserta henson �948, sample RA-0�-203, x63.

Senckenbergiana lethaea, 87 (�); 2007 Plate 2



Plate 3


Figs �, 4. Lituosepta recoarensis cati, megalospheric forms, RA-0�-�86, �, x53, 4 x43.Fig. 2. Lituosepta compressa (höttinger �967), megalospheric forms, RA-0�-�86, x60.Fig. 3. Orbitopsella praecursor (güMBel �872), RA-0�-�86, x40.Fig. 4. Lituosepta compressa (höttinger �967), megalospheric forms, RA-0�-�86, x36.Fig. 5. Amijiella amiji (henson �948), sample RA-0�-�89, x83.Fig. 6. Everticyclammina praevirguliana Fugagnoli 2000, sample RA-0�-�40, x80.


Plate 4

All thin-section hotomicrographs from Pantokrator Formation, Chalkis Quarry, Greece

Fig. �. Everticyclammina praevirguliana Fugagnoli 2000, sample CQ �07b x35.Fig. 2. Siphovalvulina gibraltarensis BouDagher-FaDel et al. 200�, sample CQ�0�, A. transver-

se section, B. Vertical section, x75.Fig. 3. A. Thaumatoporella ?parvovesiculifera (raineri �922), B. Siphovalvulina sp. sample CQ

87, vertical section, x90.Fig. 4. Riyadhella praeregularis BouDagher-FaDel et al. 200�, sample CQ�03, transverse

section, x50.Fig. 5. Everticyclammina praevirguliana Fugagnoli 2000, Sample CQ�09, x45.Fig. 6. Duotaxis metula Kristan �957, sample CQ �07b, x50.


Plate 5

Figs �-3 thin-section photomicrographs from Mt. xiron Orus, NE Evvia, Greece, Pantokrator Formation, Figs 4-6 thin section photomicrographs from Cuevas Labrados Formation, Almonacid de la Cuba, North-ern Spain.

Fig. �. Textulariopsis sinemurensis new species, sample �2.�0.6�, x83.Fig. 2. Siphophalvulina sp., sample �2.�0.64, x200.Fig. 3. Everticyclammina praevirguliana Fugagnoli 2000, sample 4.��.�, xFig. 4, 6. Bosniella oenensis gušiç, �977, 4, sample AC26, x80, 6, sample AC39, x33.Fig. 5. Palaeodasycladus ?mediterraneus (pia �920), A. Transverse section, B. oblique vertical

section. sample AC39, x�6.


Plate 6

Figs �-2 thin-section photomicrographs from Cuevas Labrados Formation, Almonacid de la Cuba, Nor-thern Spain, Figs 3-6 Thin sections from Oust Formation, Jebl Aziz, Tunisia Dorsales.

Fig. �. Cayeuxia ?piae Frollo �938, sample AC 2�, x20.Fig. 2. Palaeodasycladus ?mediterraneus (pia �920), sample AC 39, x�7.Figs 3-5. Siphovalvulina gibraltarensis BouDagher-FaDel et al. 200�, �. 3A.5 vertical sections,

sample JA37, x�00, 3B indeterminate foraminifera. 4. Transverse section, sample JA48, x�20.

Fig. 6. Siphovalvulina colomi BouDagher-FaDel et al. 200�, sample JA57, x90.


Plate 7

All thin-section photomicrographs from Jebel Rat Formation, Aghbalou n’ Kerdous, southern High Atlas, Morocco.

Fig. �. Amijiella amiji (henson, �948), sample Ad8, x36,Fig. 2. Pseudopfenderina cf. butterlini (Brun, �962), sample Ad56g, x30. Fig. 3-4. Bosniella oenensis gušiç, �977, 3, sample Ad7, x30, 4, sample Ad5, x75.Fig. 5. Cayeuxia ?piae Frollo, �938, sample Ad56c, x23.Fig. 6. Palaeodasycladus ?mediterraneus (pia, �920) entrapping A, Pseudocyclammina sp., sample Ad7, x20.


Plate 8

All thin-section photomicrographs from Jebel Rat Formation, Aghbalou n’ Kerdous, southern High Atlas, Morocco.

Fig. �. Amijiella amiji (henson, �948), sample Ad8, x89.Figs 2-5. Haurania deserta henson, �948, 3-5 sample Ad�3, 2, x 30, 3, x40, 4, x40, 5, x50. Fig. 6. Bosniella oenensis gušiç, �977, sample Ad�3, x70.


Plate 9

All thin-section photomicrographs from the Calcare Massiccio, Mt Bove south, central Apennines, Italy.

Fig. �. Textulariopsis sinemurensis new species, paratype, P.66950, sample MB�, x54.Figs 2-3. Everticyclammina praevirguliana Fugagnoli 2000, sample MB�, x60.Figs 4-5. Siphovalvulina colomi BouDagher-FaDel et al. 200�, sample MB 63, 4. Megalospheric

form showing the initial coiled part of the test, x75. 5, Microspheric form showing the nearly parallel sides in later growth, x85.

Fig. 6. Siphovalvulina gibraltarensis BouDagher-FaDel et al. 200�, sample MB30, Apennines section at Mt Bove, x�50.

Fig. 7. Thaumatoporella ?parvovesiculifera (raineri), sample MB�0, x50. Fig. 8. Thaumatoporella sp. sample MB2, x20.


Plate �0

All thin-section photomicrographs from Calcare Massiccio, Mt Bove south, central Apennines, Italy.

Fig. �. Siphovalvulina colomi BouDagher-FaDel et al. 200�, sample MB20, 0.6mm.Fig. 2. Riyadhella praeregularis BouDagher-FaDel et al. 200�, transverse section showing a

quadriserial test, sample MB22 0.25mm.Fig. 3. Cayeuxia ?piae Frollo �938, 5, sample MB 45, 2mm.Fig. 4. Thaumatoporella ?parvovesiculifera (raineri), sample MB 49, �.3mm.Fig. 5. Everticyclammina praevirguliana Fugagnoli 2000, sample MB66, �.2mm.Fig. 6. Pseudopfenderina cf. butterlini (Brun �962), a transverse section, sample MB 49,

0.2mm.

Senckenbergiana lethaea, 87 (�); 2007 Plate �0



Plate ��

All thin-section photomicrographs from the Gibraltar Limestone Formation, Gibraltar.

Figs �, 7. Siphovalvulina gibraltarensis BouDagher-FaDel et al. 200�, figured holotype P.669�2, sample G8, x, 5, figured paratype P66930, sample D20, x90.

Fig. 2. Everticyclammina praevirguliana Fugagnoli; vertical section of megalospheric form, showing an alveolar test wall and a single septal aperture, P.6694�, sample G27a. x90.

Fig. 3. Textulariopsis sinemurensis new species, 3, vertical section, holotype P66936, sample G8, 6, transverse section, x40.

Figs 4, 5. Siphovalvulina colomi BouDagher-FaDel et al., 200�, 2, vertical section, holoype P.669�0, sample G27, x�00, 4, transverse section, sample L6, paratype P669��b, x90.

Fig. 6. Riyadhella praeregularis BouDagher-FaDel et al., 200�, figured paratype, transverse section, P66947, sample D20, x68.

Figs 8, 9. Palaeodasycladus ?mediterraneus (pia). 8, PP6945, sample G5, x23, 9. P66932, G 27, x30.

Fig. �0. Thaumatoporella ?parvovesiculifera (raineri). 5, P.66949, sample E5, x40.

Senckenbergiana lethaea, 87 (�); 2007 Plate ��


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