Profit 16: 107-120, 37 Figs., 2 Tab.; Stuttgart 1999

Ammonite Biostratigraphy from the Planula and Platynota Zones in the Lugar Section (External Subbetic, Southern Spain)

J.E. CARACUEL, JAEN, F. OLORIZ& F. J. RODRI'GUEZ-TOVAR, GRANADA*

Abstract

The analysis of the precise record of ammonites in the Lugar section (Murcia, southern Spain) provides reliable data to apply the biochronostratigraphic interpretation of the Planula and Platynota Zones in ammonitico rosso and related facies outcropping in the eastern Subbetic Zone of the Betic Cordillera. The ranges of Subnebrodites planula (HEHL in Zieten, 1830), Sutneria galar (OPPEL) and Sutneria platynota (REINECKE) permit the identification of the Planula and Galar Subzones of the Planula Zone, as well as the recognition of the Platynota Zone. The subdivision of the Planula Zone in standard subzones is approached, but future research is necessary to precise correctable subdivisions within the Platynota Zone. Stratigraphic condensation and, probably, hiatuses make difficult the conclusive evaluation of the ammonite ranges in terms of the completeness of the stratigraphic record determining that zonal subdivision at Ihe level of ammonite biohorizon are limited and preliminary. Two new ammonite species, Nebrodites passendorferiiforme n. sp. and Orthosphinctes (Ardescia) betica n. sp. show potential significance for biochronostratigraphic interpretations.

1 INTRODUCTION fordian and deposits preserved in variably marly and calcareous ammonitico rosso and related facies, which are rich in ammonite re- In the Mediterranean Tethys s.str., which em- mains. In the province of Murcia, Caracuel et al. braced epioceanic environments during the Middle (1998, 1999) revisited ammonite biostratigraphy and Late , the precise ammonite biostrati- in the Lugar section in the Sierra de Lugar and pro- graphy from both the Planula and Platynota Zones, posed this section to be a reference section for the uppermost Oxfordian and/or lowermost Kimme- Oxfordian in the External Subbetic. In this section, ridgian, is limited since depositional conditions we analysed precise ammonite ranges throughout dominantly caused hiatuses in more or less con- the Planula and Platynota Zones on the basis of a densed and red nodular limestones ranging between continuous sampling of 10-20 cm thick samples marly and calcareous ammonitico rosso. Ammonite (Figs. 1-2). assemblages in Sequeiros (1974), Fulop (1976), Sapunov (1979), Sequeiros & Oloriz (1979), Sarti (1988a, 1993), Fozy (1993a, b) clearly show this 2 THE LUGAR SECTION situation. The Subbetic Zone (Betic Cordillera) and the Sierra Norte in Mallorca (Balearic Archipelago) The section investigated at the eastern part of show some of the most favourable European am- Sierra de Lugar (UTM 30SXH592309 in the topo- monitico rosso sections with ammonitiferous graphic sheet of Fortuna: 892) can be reached from Planula and/or Platynota Zones (Barthel et at., the village of Fortuna by the road between Sierra 1966; Behmel, 1970; Sequeiros, 1974; Checa & de Lugar and Sierra de La Pila (Fig. 1). Oxfordian Sequeiros, 1990; Oloriz et at., 1998, Caracuel et deposits at the Lugar section are 9.65m thick and at., 1999). In the Betic Cordillera, the eastern Ex- made of varying marly and calcareous ammonitico ternal Subbetic showing the comparatively proxi- rosso (Fig. 2). Bedding appearance in the outcrop is mal (landwards) range of epioceanic swells offers due, mainly, to diagenetic imprint, but minor dis- favourable sections with comparatively thick Ox- continuities can be recognised. The Oxfordian suc-

* Addresses of the authors: J.E. Caracuel, Depto. Ingenieria Cartografica, Geodesica y Fotogrametrfa. Escuela Politecnica Superior, Univ. Jaen, 23071 Jaen, Spain, e-mail: [email protected] Prof. Dr. Federico Oloriz Saez, Francisco Javier Rodriguez-Tovar, Depto. Estratigraffa y Paleontologi'a, Fac. Ciencias, Univ. Granada. 18002, Granada, Spain, e-mail: [email protected] -- [email protected] 108

'licanteiflpy^ W ^ < •Murcia I | Iberian im^TK a / Peninsula./ .

Mediterranean Sea

I'VO Iberian Meseta Alboran Domain •• Volcanic rocks Trough Flysch Units [V\1 External Prebetic E553 Internal Prebetic 1 Intermediate Units External Subbetic •i Middle Subbetic l-"::rl Internal Subbetic to Murcia 15Km to Murcia 15Km

Fig. 1: Geological sketch and location of the Lugar section (eastern External Subbetic). cession overlies a ferruginized hardground with Mn The Platynota section is a 1.50m thick strati- nodules ("snuff-boxes") and abundant reworked graphic interval with increasing upward carbonate ammonites and belemnites (Upper Callovian). The content (horizons 785-795 to 920-935). From uppermost Oxfordian or lowermost Kimmeridgian bottom to top, red-nodular marly limestones Planula Zone was identified within a marly am- evolving to slightly nodular rhythmic limestones monitico rosso stretch (Caracuel et al., 1999) just characterise the Platynota section. Microfacies are underlying transitional deposits from marly to cal- similar to those in the Planula Zone, although they careous ammonitico rosso and then grey and more are more bioclastic (wackestones rich in Globo- or less nodular limestones. These deposits yielded chaetes) in the upper part, where intense colonisa- the almost continuous record of the index species tion by large Thalassinoides maker's organisms Sutneria platynota (REINECKE), which permits the was widespread in some horizons. identification of the Lower Kimmeridgian Platynota Zone. Upper in the section, the remainder Kimme- ridgian and Tithonian can be recognised by ammon- 3 AMMONITE ASSEMBLAGES ites. The Planula section is made of 1.60 m of red- Horizon 640-655: Holcophylloceras mediterraneum nodular marly ammonitico rosso (horizons 640-655 (NEUMAYR), Sowerbyceras sp. A, Sowerb. sp. C, to 775-785). They are mudstones and wackestones Taramelliceras (Taramelliceras) sp. gr. costatum showing Globochaetes as dominant bioclast, along (QUENSTEDT), Taram. (Metahaploceras) wenzeli with Protoglobigerina, thin-shelled bivalves, (OPPEL), Passendorferia (Enayites) sp. gr. rozaki Saccocoma and radiolaria. Chondrites-type burrows MELENDEZ, Subnebrodites planula (HEHL in Zieten, dominate rather than Planolites although three 1830), Orthosphinctes (Praeataxioceras) sp., decimetre-thick calcareous horizons show intense Orthosph. (Orthosphinctes) sp. gr. mogosensis burrowing by Thalassinoides maker's organisms. (CHOFFAT). Except for the composition in ammonite Horizon 495-520b: Taramelliceras (Metahaplo- assemblages, no other significant difference was ceras) sp. cf. pseudowenzeii (WEGELE), Passen- recognisable in microfacies and in preservation of dorferia (Enayites) wierzbowskii MELENDEZ. macrofossils with respect to the underlying Oxfor- Horizon 680-695: Sowerbyceras sp. C, Lytoceras dian deposits. polyanchomenum (GEMMELLARO in Favre, 1876), 1 09

Fig. 2: Lugar section. Lithological column, stratigraphic distribution of selected ammonite species and biochronostratigraphy (O/K = uppermost Oxfordian or lowermost Kimmeridgian; Bimam. = Bimammatum).

Taramelliceras (Metahaploceras) sp. gr. falcula Taram. (Metahaploceras) wenzeli (OPPEL), Sut- (QUENSTEDT), Benetticeras benettii CHECA, neria galar (OPPEL). Passendorferia (Enayites) wierzbowskii MELEN- Horizon 755-770: Sowerbyceras tortisulcatum DEZ, Subnebrodites planula (HEHL in Zieten, (D'ORBIGNY), Sowerb. sp. C, Lytoceras orsinii 1830), Subneb. laxevolutum (FONTANNES), (GEMMELLARO in Favre, 1875), Taramelliceras Subneb. minutum (DIETERICH), Subneb. schroederi (Strebliticeras) sp. gr. externodosum (DORN), (WEGELE), Subneb. n. sp. A. Glochiceras (Coryceras) microdomum (OPPEL), Horizon 520-535b: Taramelliceras (Metahaplo- Sutneria galar (OPPEL), Subnebrodites laxevolu- ceras) sp. cf. kobyi quenstedti (H...LDER) - tum (FONTANNES), Subneb. minutum (DIETERICH), wegelei SCHAIRER, Clambites (Clambites) clam- Subneb. sp. gr. proteron (NITZOPOULOS), Subneb. bus (OPPEL), Pseudowaagenia micropla (OPPEL), n. sp. A, Orthosphinctes (Orthosphinctes) poiy- Passendorferia (Enayites) wierzbowskii MELEN- gyratus (REINECKE). DEZ, Sutneria sp. gr. galar (OPPEL). Horizon 775-785: Calliphylioceras manfredi (OP- Horizon 720-735: Calliphylioceras manfredi (OP- PEL), Holcophylioceras mediterraneum (NEU- PEL), Sowerbyceras sp. C, Taramelliceras MAYR), Sowerbyceras tortisulcatum (D'OR- (Strebliticeras) sp. gr. externodosum (DORN), BIGNY), Sowerb. sp. C, Lytoceras polyancho- menum (GEMMELLARO in Favre, 1876), Lyt. 110

orsinii (GEMMELLARO in Favre, 1875), Taramel- & SCHAIRER, sp. gr. binodum (OP- liceras (Metahaploceras) kobyi quenstedti PEL) - sesquinodosum (FONTANNES in Dumortier (HOLDER), Taram. (Metahaploceras) sp. gr. kobyi & Fontannes, 1876), Aspidoceras or Physodo- (CHOFFAT), Taram. (Metahaploceras) sp. aff. ceras sp., Sutneria platynota (REINECKE), Sutn. subnereus (WEGELE), Aspidoceras binodum (OP- (?) sp, Orthosphinctes (Ardescia) sp. cf. proin- PEL), Aspid. sesquinodosum (FONTANNES in conditus (WEGELE). Dumortier and Fontannes, 1876), Physodoceras Horizon 835-855: Sowerbyceras sp. A, Sowerb. aitenense (D'ORBIGNY), Benetticeras benettii sp. C, Taramelliceras (Metahaploceras) sp. aff. CHECA, Sutneria galar (OPPEL), Sutn. sp. aff. kobyi wegelei SCHAIRER, Barthelia subbetica galar (OPPEL), Subnebrodites iaxevolutum (FON- OLTRIZ & SCHAIRER, Aspidoceras sesquinodosum TANNES), Orthosphinctes (Orthosphinctes) sp. cf. (FONTANNES in Dumortier and Fontannes, 1876), polygyratus morph. colubrinus (REINECKE in Physodoceras wolfi (NEUMAYR), Sutneria platy- Oloriz, 1978). nota (REINECKE), Sutn. sp., Orthosphinctes (Ar- Horizon 785-795: Calliphylioceras manfredi (OP- descia) n. sp. aff. desmoides debelmasi ATROPS. PEL), Holcophyiloceras mediterraneum (NEU- Horizon 860-885: Aspidoceras sesquinodosum MAYR), Sowerbyceras tortisulcatum (D'OR- (FONTANNES in Dumortier and Fontannes, 1876), BIGNY), Sowerb. sp. C, Lytoceras polyancho- Physodoceras sp. cf. wolfi (NEUMAYR), Sutneria menum (GEMMELLARO in Favre, 1876), Lyt. platynota (REINECKE). orsinii (GEMMELLARO in Favre, 1875), Taramel- Horizon 880-900: Sowerbyceras silenum (FON- liceras (Metahaploceras) sp. gr. kobyi (CHOF- TANNES in Dumortier & Fontannes, 1876), Ta- FAT), Taram. (Metahaploceras) kobyi wegelei ramelliceras (Metahaploceras) sp., Physodoceras SCHAIRER, Taram. (Metahaploceras) sp. aff. sub- wolfi (NEUMAYR), Sutneria platynota (REINECKE). nereus (WEGELE), Taram. (Metahaploceras) sp., Horizon 912-917: Sowerbyceras silenum (FON- Strebiites sp. gr. tenuilobatus (OPPEL), Aspido- TANNES in Dumortier & Fontannes, 1876), Sow. ceras binodum (OPPEL), Aspid. sesquinodosum loryi (MUNIER CHALMAS in Pillet & De Fromen- (FONTANNES in Dumortier and Fontannes, 1876), tel, 1875), Lytoceras orsinii (GEMMELLARO in Physodoceras aitenense (D'ORBIGNY) morph. ai- Favre, 1875), Taramelliceras (Metahaploceras) tenense D'ORBIGNY and circumspinocum OPPEL, sp. cf. kobyi quenstedti (HOLDER in Schairer, Physod. sp. cf. aitenense (D'ORBIGNY), Benet- 1972), Taram. (Metahaploceras) sp. gr. kobyi ticeras benettii CHECA, Sutneria platynota (CHOFFAT), Taram. (Metahaploceras) sp., (REINECKE), Sutn. sp. cf. nusplingensis FISCHER, Physodoceras aitenense (D'ORBIGNY) morph. cir- Passendorferia (Enayites ?) sp., Orthosphinctes cumspinosum OPPEL, Nebrodites sp. gr. hospes (Orthosphinctes) polygyratus morph. colubrinus (NEUMAYR). (REINECKE in Oloriz, 1978). Horizon 920-935: Physodoceras sp. cf. wolfi Horizon 800-815: Sowerbyceras silenum (FON- (NEUMAYR), Nebrodites passendorferiiforme n. TANNES in Dumortier & Fontannes, 1876), Sow- sp., Sutneria platynota (REINECKE). erb. tortisulcatum (D'ORBIGNY), Sowerb. sp. C, Horizon 936-940: Taramelliceras (Metahaplo- Lytoceras orsinii (GEMMELLARO in Favre, 1875), ceras) sp. gr. strombecki (OPPEL). Lyt. polyanchomenum (GEMMELLARO in Favre, Horizon 955: Sowerbyceras silenum (FONTANNES 1876), Taramelliceras (Metahaploceras) kobyi in Dumortier & Fontannes, 1876), Sowerb. sp. wegelei SCHAIRER, Taram. (Metahaploceras) sp. C., Nebrodites sp. aff. kobyi wegelei SCHAIRER, Taram. (Metahaplo- ceras) sp. gr. kobyi (CHOFFAT), Aspidoceras bi- nodum (OPPEL), Aspid. sesquinodosum (FONTAN- 4 BIOCHRONOSTRATIGRAPHY NES in Dumortier & Fontannes, 1876), Physodo- ceras aitenense (D'ORBIGNY) morph. aitenense The biochronostratigraphic interpretation of the D'ORBIGNY and circumspinosum OPPEL, Physod. Oxfordian/Kimmeridgian boundary section, or low- sp. cf. aitenense (D'ORBIGNY), Physod. sp., Ben- ermost Kimmeridgian section, investigated at the etticeras benettii CHECA, Sutneria platynota Lugar profile is based on the composition of am- (REINECKE), Sutn. sp. cf. nusplingensis FISCHER, monite assemblages recovered from the horizons Orthosphinctes (Orthosphinctes) polygyratus quoted above. The careful palaeontological study morph. colubrinus (REINECKE in Oloriz, 1978). made on 500 ammonites complements that provided Horizon 815-825: Sowerbyceras silenum (FON- by Caracuel (1996). Present data and interpreta- TANNES in Dumortier & Fontannes, 1876), Sow- tions of the Planula Zone follow those made by Ca- erb. sp. C, Taramelliceras (Metahaploceras) sp. racuel et al. (1999), while those of the Platynota gr. kobyi (CHOFFAT) - rigidum (WEGELE), Taram. Zone improve preliminary ones provided bv Ca- (Metahaploceras) sp., Barthelia subbetica OLoRIZ racuel (1996). 1 1 1

4.1 Uppermost Oxfordian or lower- Mediterranean Tethys. The absence of Passendor- most Kimmeridgian feria (Enayites), envisaged by Caracuel et al. (1999), could be related to sampling hazard since Planula Zone the genus has been interpreted to exist, rare, in Horizons 640-655 to 775-785 (Fig. 2). The overlying horizons belonging to the Platynota Zone. lower boundary is identified through the first ap- Up resent, no very significant data for precise bio- pearance datum (FAD) of Subnebrodites planula chronostratigraphy were provided by haploceratids (HEHL) (Fig. 2; Figs. 36-37). The upper boundary because of their unfavourable preservation in am- corresponds to the last appearance datum (LAD) of monitico rosso. Notwithstanding, the range of 7a- Sutneria galar (OPPEL), just below the FAD of ramelliceras (Metahaploceras) wenzeli (OPPEL) Sutneria platynota (REINECKE) (Fig. 2; Fig. 26). from the Planula Subzone to the base of the Galar Ammonite ranges show some significant traits, Subzone is noteworthy (Fig. 2; Fig. 19) coinciding which could evidence condensation or ranges others with that notified by Wierzbowski (1978) for than usually registered from epicontinental central Poland. The known record of the Oppel' spe- deposits, such as the record of Subnebrodites to- cies in the Lugar section contrasts with those in gether with Sutn. galar (OPPEL) and no differentia- central Poland and southern as recently tion of ranges in Subnebrodites. However, even as- reported by Matyja & Wierzbowski (1997) and suming condensation, some notable biostratigraphic Schweigert & Callomon (1997), respectively. Al- data are: a) the turnover between the ranges of though the record of the Planula and Galar Subzones Sutneria sp. gr. galar (OPPEL) and Sutneria galar are probably incomplete in the Lugar section, the (OPPEL); b) the combined record of Sutneria sp. gr. over-all correlation of these two subzones with the galar (OPPEL) with Passendorferia (Enayites); c) standard subdivisions of the Planula Zone is consid- the combined record of the latter with Subnebro- ered to be acceptable. dites; d) the record of Subnebrodites above the younger Passendorferia (Enayites) wierzbowskii MELENDEZ; and e) the clearly differentiated record 4.2 Lower or lowermost Kimme- of primitive Orthosphinctes (mogosensis group) in ridgian the older horizons and conservative morphs of the polygyratus group above. Moreover, the record of 4.2.1 Platynota Zone Subnebrodites minutum within the upper range of Horizons 785-795 to 920-935 (Fig. 2). The the genus Subnebrodites is noteworthy since cor- Platynota Zone is identified through the range of the roborates recent reports from epicontinental areas index species Sutneria platynota (REINECKE) within the Betic Cordillera (Oloriz et al., 1992, and throughout the Platynota section (Fig. 2; Figs. 2 5- this volume), as well as from Poland (Matyja & 26), showing one of the thicker Platynota sections Wierzbowski, 1997). known from West Tethyan epioceanic ammonitico As noted by Caracuel et al. (1999), more re- rosso with a nearly continuous record of the index search is necessary to reach the conclusive inter- species. However, stratigraphic condensation is pretation and subdivision of the Planula Zone at the relevant comparing the known thickness of the level of ammonite biohorizons in epioceanic am- Platynota section in the close epicontinental monitico rosso. Behmel (1970) and Sequeiros & Prebetic Zone (Rodrfguez-Tovar, 1993; OI6riz & OI6riz (1979) proposed the recognition of the Galar Rodrfguez-Tovar, 1993). In absence of frequent re- Subzone in the eastern and central Subbetic, re- cords of morphologically evolved Ataxioceratinae, spectively. Condensation without doubt occurred in which support Submediterranean biochronostrati- the Lugar section, but the subdivision of the Planula graphy for the Platynota Zone in southern Europe Zone seems to be advisable even assuming peculi- (Atrops, 1982), the records of Orthosphinctes arity in some ranges of the ammonite species (Ardescia) sp. cf proinconditus (WEGELE) and Or- and/or genera registered, as assumed by Caracuel thosphinctes (Ardescia) betica n. sp. are significant et al. (1999). The Planula Subzone shows at the for biochronostratigraphy (Figs. 3-4). Thus, as- base the FAD for Subnebrodites planula (HEHL in suming the general trends of phenotypic evolution in Zieten, 1830), together with the LAD for Passen- epicontinental Ataxioceratinae assemblages, the dorferia (Enayites) group of rozaki MELENDEZ Pas- alluded records should evidence stratigraphic in- sendorferia (Enayites) wierzbowskii MELENDEZ was tervals within the middle and upper parts of the first recorded from overlying horizons of the Platynota Zone, respectively. Other interesting Planula Subzone, just below the FAD of Sutneria ammonites registered are rare Sutneria, Barthelia galar (OPPEL). The Galar Subzone is well charac- and Passendorferiinae, accompanied by more fre- terised by the record of the index species, together quent aspidoceratids and some of the so-called ido- with Orthosphinctes of Kimmeridgian affinity in the ceratids. 1 1 2

Registered between the upper Planula and the Platynota Zones, but mainly within the latter, there 5 REMARKS ON AMMONITES are transient forms between very evolute specimens interpreted as Nebrodites, Mesosimo- The following comments are restricted to the ceras, and Passendorferia (Brochwicz-Lewinski & new ammonite species registered: Orthosphinctes Rozak, 1976), "Passendorferia" s.l. (Sarti, (Ardescia) betica n. sp. and Nebrodites 1990), and several genera of Idoceratidae (Sarti, 1993). Ammonites belonging to this group were re- passendorferiiforme n. sp. ported from the Platynota Zone in the central Sub- Superfamily Perisphinctaceae Steinmann betic Zone by OI6riz (1978), but their record is 1 890 better known from northern (Pavia et al., 1987; Sarti, 1993). In the Lugar section they are Family Ataxioceratidae Buckman 1921 represented by rare Passendorferia (Enayites), Ne- brodites passendorferiiforme n. sp. (Fig.4a-b), and Subfamily Ataxioceratinae Buckman 1921 fragmented material difficult to identify at the ge- nus/species level. In general, precise biostratigra- Genus Orthosphinctes Schindewolf 1925 phy in this group of very evolute ammonites is lim- ited and, therefore, their phylogenetic relationships Subgenus Ardescia Atrops 1982 are poorly known showing a rather intricate pat- tern of evolution between Passendorferiinae and the Type species Ataxioceras desmoides Wegele 1929 so-called idoceratids. Thus, the phylogenetic rela- tionships that Sarti (1990) envisaged appear to be Orthosphinctes (Ardescia) betica n. sp. a valuable but oversimplified interpretation. Tb. 1, Figs. 3-4 Among aspidoceratids, Benetticeras benettii Synonymy: 1996 Orthosphinctes (Orthosphinctes) sp. CHECA is frequent in the lower part of the C - CARACUEL, p. 249 Platynota Zone, although it is known from the mid- dle-upper Planula Subzone in the section. However, Holotype: LG-855-1 data from the central Subbetic Zone indicate its re- Locus tipicus: Sierra de Lugar, Province of Murcia, cord from the entire Planula and Platynota Zones in Spain. the Betic Cordillera (Checa & Oloriz, 1984; Checa, Stratum tipicum: Horizon 835-855 at the Lugar 1985). section, Lower Kimmeridgian, upper Platynota A similar case affects to the rare Barthelia sub- Zone. betica OLORIZ & SCHAIRER, which was previously Derivatio nominis: In allusion to the region of known to range from the Bimammatum to the provenance, the Betic Cordillera. Planula Zones in the Betic Cordillera (OI6riz & Material: one specimen Schairer, 1983; Oloriz et al., 1992, and this vol- ume). Its record from the middle and the lower up- per part of the Platynota Zone in the Lugar section could evidence size reduction upwards according to the youngest specimen registered (Fig. 2; Figs. 16- 17). Although this observation cannot be conclusive, the youngest record suggests the revi- sion of phylogenetic interpretations as envisaged by Schweigert (1995). The range of Sutneria sp. cf. nusplingensis FISCHER characterises a lower part of the Platynota section investigated (Fig. 2, Figs. 2 2- 24). The range registered fits well with obser- vations made by Geyer (1961) who assumed Jhe record of galar-type Sutneria in the lowermost part of the "Badenerschichten" in the "Aargauer Jura" of northern Switzerland. This species is also known from equivalent horizons in the Prebetic Zone (Oloriz & Rodriguez-Tovar, 1996). Figs. 3-4: Orthosphinctes (Ardescia) betica n. sp. LG-855- 1. Ventral and left-side views of the holotype. Horizon LG-855. x1. 1 1 3

Tab. 1: Shell measurement3me s (mm) of specimen LG-855-1, holotype of Orthosphinctes (Ardescia) betica n. sp.

Dm \ U W H UR U/Dm W/Dm H/Dm W/H 63 \29 15 1 9 54 0,46 0,24 0,30 0,79 56 25 1 4 1 8 52 0,45 0,25 0,32 0,78 52 21 1 8 51 0,40 0,23 0,35 0,67 39 17 11 14 49-50 0,44 0,28 0,36 0,79 35 1 5 11 14 47 0,43 0,31 0,40 0,79

Diagnosis: Small size. Bifurcate, inter- Ardescia shows species with similar sculpture at calatory and polygyrate ribs. Constrictions subtle the upper Desmoides and Guilherandense Subzones or absent in the phragmocone, but large and shallow of the Platynota Zone (see Atrops, 1982), but rib on the body chamber. Parabolae absent. Peristome density is higher in specimen LG-855-1 and the unpreserved. reverse affects to ribbing index in the French Species. Species from the middle-upper Platynota Description: Incomplete specimen of Zone showing closer rib curves, such as 63mm in diameter showing the phragmocone and the Orthosphinctes (Ardescia) tintanti ATROPS, rear quarter of the body chamber preserved. Orthosph. (Ardescia) proinconditus ATROPS, and Coiling is moderate (U/Dm=40-46%). In the outer Orthosph. (Ardescia) n. sp. A ATROPS, are greater whorl, the umbilicus (U=29mm) shows a small, in size, more evolute, and show less dense and vertical umbilical wall. Although whorl-width stronger ribbing. Morphs of Orthosph. (Ardescia) values in the body chamber are only approximate schaireri ATROPS exist showing similar ribbing (W/Dm=24%) crushing is negligible. The whorl index, but they are less dense ribbed and usually section is ovate (W/H=0,67-0,79) with slightly developed constrictions and typical parabolic convex flanks and relatively narrow ventral region. structures. Orthosph. (Ardescia) desmoides Maximum whorl width is located very close of the desmoides (WEGELE) and Ataxioceras (Schneidia) prominent umbilical edge. Ribbing is dominated by guilherandense ATROPS (the latter rarely bifurcates on the phragmocone, with occasional developed ataxioceratid ribs) show rib curves intercalation of external ribs (one between two decreasing from 20-35mm and 35-50mm, bifurcates) and polygyrates on the last whorl. The respectively, developed constrictions frequently, beginning of the body chamber shows a sharp and the latter experienced a variable trend to decrease in density of primary ribs, accompanied reinforced primaries on the umbilical shoulder. by dominant polygyrate ribs with additional Orthosph. (Ardescia) desmoides debelmasi ATROPS, intercalatories as in the phragmocone. Close to the the species of subgenus Ardescia closest to Atax. end of the preserved body chamber there is a large, (Schneidia) guilherandense ATROPS, can be similar shallow and prorsiradiate constriction with in sculpture as well as coiling, but its ribbing curve projected sinus on the mid-venter. decreases from 40mm onwards. Orthosph. (Ardescia) thieuloyi ATROPS is smaller, and typically shows constrictions and rib curve Re m a r k s : The specimen LG-855-1 shows a decreasing from 30mm. primitive rib curve (at 35, 39, 52, 56 and 63mm shows 47, 49-50, 51, 52 and 54 primary ribs) Since the group of desmoides WEGELE, even with positive but lower slope than in epicontinental rare, is known from the Subbetic Zone (Oloriz, Orthosphinctes. The ribbing index (number of 1978), while the presence of Ataxioceras external ribs per 10 umbilical ribs) on the (Schneidia) cannot be proved from the region, and phragmocone varies between 2.2 at 37mm and 2.7 ataxioceratid ribs are absent from specimen LG- at 54mm. Orthosphinctes (Orthosphinctes) 855-1, we prefer to interpret this specimen as poiygyratus (REINECKE) shows similar shell Orthosphinctes (Ardescia). Hence, Orthosphinctes morphology and sculpture in shells that typically (Ardescia) betica n. sp. exemplifies a case of are greater in size, and the range of this species in limited and probably diachronic sharing of the main epicontinental areas is shorter and older. trends in morphologic evolution in epioceanic and Mediterranean Orthosphinctes developed a more epicontinental ataxioceratins during the middle to conservative ribbing style (Oloriz, 1978; Sarti, late Platynota Chron. 1993). Among the morphologically more evolved epicontinental Orthosphinctes, the subgenus 1 1 4

Family Perisphinctidae Steinmann 1890 Locus tipicus: Sierra de Lugar, Province of Murcia, Spain. Subfamily Passendorferiinae Melendez Stratum tipicum: Horizon LG-920-935 at the Lugar 1 989 section, Lower Kimmeridgian, upper Platynota Zone. Genus Nebrodites Burckhardt 1912 Derivgtio nominis: In allusion to the very evolute, Passendorferia-like, type of shell. Type species Simoceras agrigentinum Gemmellaro Material: Two specimens and one fragment. 1872 Diagnosis: Small size. Very evolute. Ribs Nebrodites passendorferiiforme n. sp. simple and bifurcate. Prorsiradiate constrictions in Tab. 2, Figs. 5-14 phragmocone and body chamber. Peristome unpreserved. Holotype: LG-920-935-5

Figs. 5-9: Nebrodites passendorferiiforme n. sp. LG-920-935-5. Right-side and ventral views of outer and inner whorls of the holotype. Horizon LG-920-935. x1.

Figs. 10-11: Nebrodites passendorferiiforme n. sp. Right-side and ventral views of small fragment of body chamber belonging to a specimen slightly greater than the holotype. Horizon LG-920-935. x1.

Figs. 12-14: Nebrodites passendorferiiforme n. sp. LG-920-935-39. Right-side and ventral views of phragmocone and right-side view of the same specimen with incomplete body chamber. Horizon LG-920-935. x1.

Tab. 2: Shell measurements (mm) of specimens LG-920-935-5 (holotype) and LG-920-935-39 (paratype) of Nebrodites passendorferiiforme n. sp. [*] approximate.

Dm U W H UR U/Dm W/Dm H/Dm E/H 30 1 8 6.5 7 49 0,60 0,21 0,23 0,92 24,5 14,5 6,2 5,5 46 0,59 0,25 0,22 1,12 19,5 1 1,2 5,9 4,5 42 0,57 0,30 0,20 1,31 14,9 8,4 3,2 34 0,56 0,21

LG- 920-935- •39 Dm U W H UR U/Dm W/Dm H/Dm E/H 32 1 8 6 9 4 1 * 0,56 0,19 0,28 0,67 29 1 6 6 8 40* 0,55 0,21 0,28 0,75 24 1 4 5 6 36' 0,58 0,21 0,^5 0,83 115

Description: Two incomplete specimens, No significant differences in shell parameters LG-920-935-5 (holotype) and LG-920-935-39 have been identified in the paratype (LG-920-935- (paratype), and one fragment of body chamber (LG- 39), but ribbing density is lower. According to 920-935-2). The holotype is 30mm in size, and body-chamber fragment LG-920-935-2, the adoral preserves the phragmocone and almost the rear projection of secondary ribs increases closer to the quarter of the body chamber. Coiling is very loose, peristome, and/or in slightly greater specimens. decreasing during the ontogeny (U/Dm=56% to Although preservation impedes the precise ob- 60%). The umbilicus is large (U=8,4 at 14,9mm; servation of suture lines, the absence of significant 11.2 at 19.5mm in Dm; 14.5mm at 24.5mm in Dm; crowding in the holotype indicates that the final 18mm at 30mm in Dm), shallow, with a very small growth was not reached, but a slightly reduced and vertical umbilical wall in the inner whorls, separation of sutures at the end of the phragmocone which even decreases in the body chamber. Whorl- could indicates that maturity should be close. In the width values are higher in the phragmocone and de- paratype, the relative crowding of suture lines in- crease during the ontogeny (W/Dm=25% to 30%) dicates an adult specimen with half-whorl pre- to be clearly lower in the body chamber served body chamber at ca. 35mm. Therefore, the (W/D=21.6%). The whorl section is depressed on new species is considered to be microconchiate be- the inner whorls (E/H=1.31), the depressed- longing to the Benacoceras-Trenerites group. subrrounded (E/H=1.12), and ovate in the body chamber (E/H=0.92). The flanks are slightly con- Remarks: The description above is mainly vex, subparallel in the phragmocone and convergent based on the holotype, which is the best-preserved to the periphery in the body chamber. The venter is specimen among these macerrimus-like ammonites. large and flat in the camerate shell, and reduces as In fact, these ammonites were interpreted as Ne- consequence of whorl-section evolution, especially brodites (Nebrodites) macerrimus (QUENSTEDT) in in the body chamber. Maximum whorl width is lo- Caracuel (1996), assuming the range of the cated close to the mid-flank in the phragmocone, Quenstedt' species to be slightly longer (slightly and comes down in body-chamber flanks. earlier FAD) that usually admitted. Later, a rare The sculpture is made of constrictions and ribs, record of Nebrodites (Nebrodites) aff. macerrimus without parabolic structures. Four to five deep (QUENSTEDT) was reported from the upper constrictions per whorl strangulate flanks and Platynota Zone (Guilherandense Subzone) in north- venter, but not the dorsal shell, from the nucleus to ern Africa. (Benzaggagh & Atrops, 1997). the last preserved whorl; the last constriction The stratigraphic interpretation of Ammonites observed belongs to the rear part of the body macerrimus QUENSTEDT has been based on Ziegler chamber. The constrictions are parallel to ribbing, (1959) who determined its age as "Oberen Malm y showing prominent adoral collar and prorsiradiate (vielleicht auch im unteren Malm 8)". This author sinus on the mid-venter. assumed that the "Weissjura a, Lochen" inter- Ribs are simple and bifurcate, without major preted by Quenstedt (1887-1888) for his Ammon- differences in relative rib density neither during ites macerrimus was a confusion, and his conclu- the ontogeny nor between the phragmocone and the sions influenced later stratigraphic interpretations preserved body chamber. Two, and rarely three, (Geyssant, 1966; Oloriz, 1978; Sapunov, 1979; consecutive simple ribs sporadically disturb the Benzaggagh, 1988; Benzaggagh & Atrops, 1997). alternation of simples and bifurcate. Ribs are radial Hence, and with rare exceptions based on scanty on the phragmocone, with divisions located on the records, the admitted range of this species resulted ventral margin determining short-ventral seconda- to be middle-late Early to early-middle Late ries and, therefore, secondary ribs are hidden in Kimmeridgian in age (Ziegler, 1959; Geyssant, the umbilicus. Towards the end of the phragmocone 1966; OI6riz, 1978; Sapunov, 1979; Benzaggagh, and the preserved body chamber, ribs are slightly 1988; Benzaggagh & Atrops, 1997), although some convex adorally and prorsiradiate, showing lower of the depicted material (Sapunov, 1979) should be divisions on the upper flanks. External ribs are not co-specific. However, a re-interpretation of typically interrupted on the mid-venter, with occa- the bio-horizon of macerrimus QUENSTEDT has been sional exceptions of slightly convex ribs. In the made by Schweigert & Callomon (1997), who re- phragmocone, ventral ribs are radial and show mid- ported this species from the Upper Oxfordian (bi- line smoothing and then a narrow smooth band, mammatum-borizon) in the type-area of Lochen in which enlarges towards the body chamber, inter- southern Germany, from which Quenstedt (1887- rupt them. On the preserved body chamber, ribs 1888) reported specimens of Epipeitoceras in his are clearly projected adorally on the venter, and "Weissjura a" (and "Weissjura p" for other ar- the enlarged ventral smooth-band separates the eas). On this basis, a new case of strong phenotypic sculpture from both shell sides. 116

convergence is demonstrated in Upper Jurassic in rib density are identifiable, but the Quenstedt' ammonites. species developed a reverse trend with decreasing According to the above, the macerrimus-like density of ribs on the outer whorl. Although re- ammonites recovered from the uppermost ported without illustration, Nebrodites (Nebrodites) Platynota Zone (Lower Kimmeridgian) in the Lugar aff. macerimus (QUENSTEDT) in Benzaggagh & section are interpreted as Nebrodites passendor- Atrops (1997) is envisaged as a potential record of feriiforme n. sp. The new species differs from the Nebrodites passendorferiiforme n. sp. in northern type of Quenstedt (plaster cast) in more frequent Africa (Prerif). Younger records of macerrimus- constrictions, less rursiradiate ribbing on the outer like ammonites in northern Africa, as well as in whorl, more external divisions of ribs on umbilical Europe, need of careful revision based on precise whorls hiding secondary ribs in the umbilicus, and a stratigraphy before to be conclusively reinter- comparatively delayed change from subrrounded to preted. ovate whorl-section. In addition, minor differences

Specimen depicted at natural size

Fig. 15: Aspidoceras sesquinodosum (FONTANNES in Dumortier and Fontannes, 1876). LG-785-815-42. Left-side view. Horizon LG-775-785.

Figs. 16-17: Barthelia subbetica OLtRIZ & SCHAIRER. LG-835-855-12. Left-side and ventral views, showing pre-peristomal constriction. Horizon LG-835-855.

Fig. 18: Physodoceras altenense (D'ORBIGNY) morph. aitenense D'ORBIGNY. LG-755-775-21. Left-side view. Horizon LG- 775-785.

Fig. 19: Taramelliceras (Metahaploceras) wenzeli (OPPEL). LG-640-655-13. Right-side view. Horizon LG-640-655.

Figs. 20-21: Taramelliceras (Metahaploceras) kobyi wegelei SCHAIRER. LG-785-795-2. Ventral and left-side views. Horizon LG-785-795.

Figs. 22-24: Sutneria sp. cf. nusplingensis FISCHER. LG-785-815-4. Left and ventral views. Horizon LG-800-815.

Fig. 25: Sutneria platynota (REINECKE). LG-920-935-4. Right-side view. Horizon LG-920-935.

Fig. 26: Sutneria platynota (REINECKE). LG-785-2. Left-side view. Horizon LG-785-795.

Figs. 27-28: Sutneria galar (OPPEL). LG-775-785-89. Right and ventral views. Horizon LG-775-785.

Fig. 29: Sutneria galar (OPPEL). LG-785-815-10. Left-side view of the body chamber. Horizon LG-775-785.

Figs. 30-31: Orthosphinctes (Orthosphinctes) polygyratus morph. colubrinus (REINECKE in Oloriz, 1978). LG-785-815-7. Right-side and ventral views. Horizon LG-800-815.

Figs. 32-33: Subnebrodites schroederi (WEGELE). LG-680-695-3. Ventral and left-side views. Horizon LG-680-695.

Figs. 34-35: Orthosphinctes (Ardescia) sp. cf. proinconditus (WEGELE). LG-825-3. Ventral view of inner whorls showing ventral parabolae, and left-side view including outer whorls with incomplete body chamber. Horizon LG-815-825.

Fig. 36: Subnebrodites planula (HEHL in Zieten, 1830). LG-640-655-11. Left-side view of a large phragmocone. Horizon LG-640-665.

Fig. 37: Subnebrodites planula (HEHL in Zieten, 1830). LG-680-695. Right-side view of juvenile showing the rear part of the body chamber. Horizon LG-680-695.

J 1 1 7 1 18

The group of hospes NEUMAYR has been by long parabolae, and does not show such an adoral pro- time considered to be the earliest record of small jection of concave ribs. Concerning to the latter, Kimmeridgian Nebrodites. The oldest records of this the only closer species idoceroides DORN is clearly group have been reported from the Platynota Zone. more strongly coiled and shows a different ribbing. They were interpreted to belong to the lowermost Among other species from the Bimammatum Zone Platynota Zone (Orthosphinctes Subzone) in the without parabolae, rozaki MELENDEZ in Atrops & Iberian Chain and the Prebetic Zone in Spain (Atrops Benest (1986, pi. 1, figs. 7-8) has radial ribbing & Mel6ndez, 1984; Rodrfguez-Tovar, 1993), as and subrrounded-depressed whorl section. From the well as in the Algarve in southern (Ro- Planula Zone, Enayites wierzbowskii (Wierzbowski, dri'guez-Tovar, 1993; Marques et al., 1998). In 1978, pi. 2, fig. 17; Matyja & Wierzbowski, 1997, addition, they were reported from the upper pi. 9, fig. 2) is greater in size, less evolute, and Platynota Zone (Guilherandense Subzone) in the In- shows radial ribbing and parabolae. ternal Prerif (Benzaggagh, 1988; Benzaggagh & More research is necessary before to elucidate Atrops, 1997). According to usual interpretations, the significance of transitional forms between Pas- the group of Nebrodites hospes shows a notable sendorferiinae showing Oxfordian and Kimmeridgian variability in coiling, constrictions and ribbing affinity. These ammonites are rare and known from (OI6riz, 1978, p.171; Rodrfguez-Tovar, 1993, the Tethyan realm and surrounding areas. In the p.180; Sarti, 1993, p.96), but its morphs closer to Prebetic Zone of the Betic Cordillera in southern macerrimus-Wke ammonites are greater in size and Spain, some of these rare ammonites, interpreted with coarser ribbing, hence clearly separated from as Idoceratinae gen. ind. and Nebrodites sp. in Nebrodites passendorferiiforme n. sp. Garci'a-Hernandez et al. (1979), and as Nebrodites The younger Nebrodites hospes hospes (NEU- sp.1 aff. hospes hospes (NEUMAYR) in OI6riz & MAYR) depicted by Sarti (1993, pi. 15, fig. 2) Rodrfguez-Tovar (1993), deserve especial at- from the lower Herbichi Zone (Stenonis Subzone) is tention. slightly more involute than Nebrodites passendor- feriiforme n. sp. Sarti's specimen shows numerous constrictions and sculpture on the outer whorl, 6 CONCLUSIONS which could indicate looser affinity with the Neu- mayr' species, as least according to the type and The Lugar section shows favourable conditions to investigate ammonite biostratigraphy in the its closer morphs. In contrast, the specimen recov- Planula and Platynota Zones in the eastern Subbetic ered from the Trento Plateau seems to be morpho- Zone of the Betic Cordillera, southern Spain. The logically closer to a phyletic line connecting Nebro- ranges of the index species Subnebrodites planula dites passendorferiiforme n. sp., and its relatives, (HEHL in Zieten, 1830), Sutneria galar (OPPEL) and with the group of Nebrodites envisaged as the an- Sutneria platynota (REINECKE) permit the iden- cestors of Mesosimoceras by Sarti (1988b). The tification of the Planula and Galar Subzones of the new species described clearly separates from these Planula Zone, as well as the recognition of the younger Nebrodites due to the persistently radial Platynota Zone. At present, the subdivision of the course of ribbing and rounded-depressed whorl Platynota Zone in the Lugar section is preliminary, sections in the latter. but a lower, middle, and upper parts of the Among the new material recently illustrated Platynota Zone seems to be identifiable by the first from the Platynota Zone, Idoceras (Subnebrodites) time in ammonitico rosso and related facies in sp. (Schairer & Schlampp, 1995, pl.1, figs. 1,2, 4) South Spain. However, stratigraphic condensation is more involute, has stronger ribbing and could be and, probably, hiatuses make difficult a conclusive morphologically close to juvenile Trenerites ac- evaluation of the ammonite ranges in terms of the cording to illustrations in Sarti (1993, pi. 22). Ido- completeness of the stratigraphic record. Zonal ceras (Subnebrodites) sp. (Gradl & Schairer, 1997, subdivision at the level of ammonite biohorizon is pi. 1, fig. 3) shows rursiradiate ribs from the limited and preliminary. The two new ammonite nucleus, less dense ribbing on the outer whorl, and species recognised, Nebrodites passendorferiiforme different whorl section (subrrounded) and con- n. sp. and Orthosphinctes (Ardescia) betica n. sp. strictions. The paratype of Trenerites enayi SARTI show potential significance for biochronostrati- (Sarti, 1993, pi. 22, fig. 2b-c) shows inner whorls graphic interpretations. with radial constrictions, while the inner whorls of the holotype (Sarti, 1993, pi. 21, fig. 1a) show a more distant sculpture. ACKNOWLEDGMENTS Among microconchiate Passendorferiinae from the Bimammatum and Planula Zones, Passendorferia This research was made within the program and (Enayites) is greater in size, frequently has financial support of the EMMI Group (RNM-178

J 119

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