Palaeodiversity 1: 133–140; Stuttgart, 30.12.2008. 133

Juraphyllites mimatensis (d’Or b i g n y , 1845) (Ammonoidea: Phylloceratida), a Tethyan immigrant in the Upper Pliensbachian of Franconia (Jurassic, Southern Germany)

He l m u t Ke u p p & Gü n t e r Sc h w e i g e r t

Abstract Two well-preserved small-sized specimens of phylloceratoid ammonites are reported from the Late Pliensba- chian Spinatum Zone of Franconia. They are assigned to Juraphyllites (cf.) mimatensis (d’Or b i g n y , 1845) and re- garded as Tethyan immigrants. Due to their very similar early ontogenetic stages it seems impossible to distinguish tiny juveniles of Juraphyllites mimatensis (d’Or b i g n y , 1845) from those of contemporaneous Meneghiniceras exi- mium (v. Ha u e r ). Keywords: Phylloceratina, Early Jurassic, palaeobiogeography, immigration, Franconia, Germany.

Zusammenfassung Zwei gut erhaltene kleinwüchsige Exemplare phylloceratoider Ammoniten werden aus der Spinatum-Zone des Ober-Pliensbachium von Franken beschrieben. Sie werden zu Juraphyllites (cf.) mimatensis (d’Or b i g n y , 1845) ge­ stellt und als tethyale Einwanderer angesehen. Wegen ihrer sehr ähnlichen frühontogenetischen Stadien erscheint es praktisch unmöglich, sehr kleine Individuen von Juraphyllites mimatensis (d’Or b i g n y , 1845) von solchen des gleich­zeitig vorkommenden Meneghiniceras eximium (v. Ha u e r ) zu unterscheiden.

1. Introduction 2. Provenance of the studied material

The dark claystones of the Upper Pliensbachian The Middle Liassic Amaltheenton Formation (Upper Amaltheenton Formation of Southern Germany, particu- Pliensbachian) of Franconia crops out in the western fore- larly of Franconia, yield a relatively low diverse ammonite land of the Northern and Middle Franconian Alb and is fauna, generally dominated by the Subboreal family of the exposed in several clay-pits. Today most of them are aban- Amaltheidae. Rare immigrations of various Tethyan or doned (e. g., Kalchreuth: Ri c h t e r 1993; Marloffstein: Euroboreal ammonite taxa seem to be restricted to short Pl ü c k e b a u m 1985; Sc h m i d t -Ka l e r 2004; Reichenschwand: phases of sea level highstands, as recently pointed out by Ha a r l ä n d e r 1961; Schnaittach: Vo i g t 1968; Unterstür- Sc h w e i g e r t (2005) and Ke u p p & Sc ho b e r t (2008). These mig: Ri c h t e r 1977, 2000; Pl ü c k e b a u m 1985). The only exotic ammonites attracted special attention and were not- still active clay-pit of the Liapor Company in this area lies ed by numerous former authors (see Sc h w e i g e r t 2005 for in the Oberfranken district, about 1 km SSE of Butten- references; Ma i s c h 2007). heim. This clay-pit substitutes the former clay-pit of Un- Soon after the publication of some findings of juvenile terstürmig which was located only 1.5 km to the south. phylloceratids from the clay-pit of Buttenheim in Franco- The latter outcrop became popular by the occurrence of nia (Ke u p p & Sc ho b e r t 2008), two somewhat larger speci- well-preserved white-coloured amaltheids bearing their mens were found by the amateur collector Joh a n n Sc ho ­ original aragonitic shells (Du l l o 1981). The stratigraphic b e r t (Hirschaid). Both specimens exhibit diameters of 30 sections exposed in both clay-pits are almost identical and 43 mm, respectively, and are assigned to the unkeeled (Ri c h t e r 2003; Ho f f m a n n et al. 2007). In the former clay- juraphyllitid taxon Juraphyllites (cf.) mimatensis pit of Unterstürmig especially the upper part of the Spina- (d’Or b i g n y , 1845). This taxon, which is of Tethyan origin, tum Zone of the Upper Pliensbachian and the Lower Toar- was previously unknown from the Pliensbachian of South- cian was exposed; only from time to time expanding ern Germany. The new findings, which question earlier downwards to the Apyrenum Subzone of the Upper Pliens- determinations of juveniles, stimulated this study. bachian. In contrast, the clay-pit near Buttenheim exposes primarily beds of the Gibbosus and Apyrenum subzones, Acknowledgements expanding upwards during the progress of the last two Joh a n n Sc ho b e r t (Hirschaid) is thanked for his generous donation of his interesting findings for this study. The journals’ years to the upper part of the Spinatum Zone and the base referees Jo a c h i m Bl a u (Gießen) and Ch r i s t i a n Me i s t e r (Genève) of the Lower Toarcian Posidonia Shale (Fig. 1). Also in the are thanked for their advice and suggestions. clay-pit of Buttenheim well-preserved ammonites occur 134 p a l a e o d i v e r s i t y 1, 2008

Fig. 1. Geographical position und section of the clay pit near Buttenheim. The horizon where the Juraphyllites mimatensis (d’Or b i g n y , 1845) illustrated in Fig. 3 was found is marked by an asterisk. k e u p p & s c h w e i g e r t , juraphyllites m i m a t e n s i s i n so u t h e r n g e r m a n y 135 with their primary aragonitic shells, sometimes, especially Remarks. – In several recent publications dealing within beds of the Apyrenum Subzone, showing a colour with Juraphyllitidae, the genera Meneghiniceras Hy a t t , pattern formed by dark, sub-concentric dotted spiral lirae 1900, Harpophylloceras Sp a t h , 1927, and Juraphyllites (Ri c h t e r 2003; Ke u p p 2005; Ke u p p & Sc ho b e r t 2008). Mü l l e r , 1939 were considered to be synonymous or of Within a collection of ca. 17.000 ammonites from the subgeneric rank (e. g., Me i s t e r 1989; Ma c c h i o n i 2001; Upper Pliensbachian of Buttenheim there are only less than Ma c c h i o n i in Pa v i a & Cr e s t a 2002; Ma c c h i o n i & Me i s t e r ten phylloceratids, most of them juveniles. The first phyl- 2003). Indeed, Harpophylloceras can be easily included in loceratid specimen described in this study was originally Meneghiniceras (Sc h w e i g e r t 2005), but the latter differs included in a small calcareous concretion. It was collected from other Juraphyllitidae by the presence of a keel in the by J. Sc ho b e r t (Hirschaid) in early March 2008 in the adult stage. Therefore we maintain the genus Juraphyllites higher section of the clay-pit of Buttenheim. It came from for the forms lacking a keel. undisturbed sediments 20 cm below the main fossiliferous horizon of the upper Spinatum Zone (= Hawskerense Sub- zone). The predominant ammonites of this level are the Juraphyllites mimatensis (d’Or b i g n y , 1845) different morphotypes of Pleuroceras spinatum (Br u ­ Figs. 2–3, ? 4 g u i è r e ) which occur mainly in the directly overlying beds. * 1845 Ammonites Mimatensis, d’Or b i g n y , 1844. – c ho The second specimen was found loose also by J. S ­ d’Or b i g n y , p. 344, pl. 110, figs. 4–6. b e r t , together with dispersedly distributed amaltheids of 1856 Ammonites Mimatensis, d’Or b ., 1845. – Op p e l , the Apyrenum Subzone in early spring 2007. p. 252. The studied ammonites are housed in the Institute of 1929 Rhacophyllites spec. ex aff. mimatensis Or b . – Gu g e n b e r g e r , p. 255, pl. 10, fig. 5, pl. 11, fig. 6, Geological Sciences, Freie Universität Berlin, collection pl.12, fig. 10. Ke u p p , numbers MAn-3111 and MAn-3115. ? 1930 Rhacophyllites cf. mimatensis (d’Or b i g n y ). – Mo n e s t i e r , p. 8, pl. 11, fig. 1. 1934 Rhacophyllites mimatensis (d’Or b i g n y ). – Mo n e s ­ 3. Systematic palaeontology t i e r , p. 16, pl. 5, figs. 17, 19, 24–30, 34–35. 1934 Rhacophyllites libertus (Ge m m e l l a r o ). – Mo n e s ­ t i e r , p. 17, pl. 6, figs. 1, 6. Suborder Phylloceratina Ar k e l l , 1950 ? 1934 Rhacophyllites mimatensis (d’Or b i g n y ) var. vin- Superfamily Phylloceratoidea Zi t t e l , 1884 ensis. – Mo n e s t i e r , p. 17, pl. 5, figs. 36–37. Family Juraphyllitidae Ar k e l l , 1950 pars 1963 Rhacophyllites mimatensis (d’Or b i g n y ). – Sg u a z ­ zo n i , p. 46, pl. 21, fig. 10, non fig. 4. ? 1970 Juraphyllites libertus (Ge m m e l l a r o ). – Br a m b i l ­ Genus Juraphyllites Mü l l e r , 1939 l a , p. 262, pl. 55, fig. 2. 1977 Juraphyllites mimatensis (d’Or b i g n y ). – Wi e d e n ­ Type species: Phylloceras diopsis Ge m m e l l a r o , 1884. m a y e r , p. 36, pl. 1, figs. 3, 8b.

Fig. 2. Juraphyllites mimatensis (d’Or b i g n y , 1845), holotype, lost; reproduction of d’Or b i g n y ’s illustration; Upper Pliensbachian; Mt. Mimat, Dept. Lozère, France. Diameter 42 mm. 136 p a l a e o d i v e r s i t y 1, 2008

Fig. 3. Juraphyllites mimatensis (d’Or b i g n y , 1845); Pliensbachian, upper part of the Spinatum Zone, Hawskerense Subzone; But- tenheim, northern Bavaria, Germany; MAn-3111. 29.2 mm in diameter. – A. Lateral view. B. Ventral view. C. Oblique view showing the large missing shell fragment due to a presumed crustacean attack. D. Close-up of Fig. 3A showing the suture line.

1989 Juraphyllites mimatensis (d’Or b i g n y ). – Me i s t e r , France and named after the type locality, the Mont Mimat south p. 30, pl. 2, fig. 10, pl. 3, figs. 4, 17a, b. of Mende (Dept. Lozère). Because of the apparent lost of the 1994 Juraphyllites mimatensis (d’Or b i g n y ). – Jo l y in type specimen originally housed in the Re n a u x collection in Fi s c h e r , p. 98, fig. 31, pl. 27, figs. 3a–c, 4. Montpellier, Jo l y (in Fi s c h e r 1994) designated a neotype, a 2000 Juraphyllites mimatensis (d’Or b i g n y , 1845). – mould coming from the middle Upper Pliensbachian (Margari- Jo l y , p. 29–30, figs. 41–43, pl. 3, fig. 6a, b. tatus Zone) of Mt. Guilhaumard near Tournadous. This neotype, however, is much less complete than the lost holotype, following Types: Juraphyllites mimatensis was first described by the illustration by d’Or b i g n y (1845) (Fig. 2). d‘Or b i g n y (1865) from the Upper Pliensbachian of Southern Occur rence: From different localities of the Grands Caus­ k e u p p & s c h w e i g e r t , juraphyllites m i m a t e n s i s i n so u t h e r n g e r m a n y 137

Tab. 1. Measurements of specimens related to Juraphyllites mimatensis (d = diameter; u = umbilicus; wh = whorl height; ww = whorl width).

d u u/d wh wh/d ww ww/d ww/wh specimen (mm) (mm) (%) (mm) (%) (mm) (%) (%) MAn-3111 29.2 (Fig. 3) 24.2 6.6 27 11.6 48 7.9 32.5 68 MAn-3115 43 6.8 15.8 24.0 56 12.1 28 50 (Fig. 4) Neotype 34.5 10 29 16 46 9.8 28 61 (Jo l y in Fi s c h e r 1994) Coll. Jo l y no. 99-1 Cornus 27.3 8.8 32 12.3 45 7.3 27 59 (Jo l y 2000) Coll. Jo l y no. 140-1 Puech 22 6.8 31 9.7 44 6 27 62 (Jo l y 2000)

ses (Southern France, e. g., d’Or b i g n y 1845; Op p e l 1856; Mo n e s ­ On the body chamber simple ribs occur, running t i e r 1934; Me i s t e r 1989; Jo l y in Fi s c h e r 1994, Jo l y 2000), slightly convex over the flattened flanks, and disappear on ­Turkey (Gu g e n b e r g e r 1929); ? Morocco (Mo n e s t i e r 1930), ? the rounded venter. Immediately behind the peristome, the West-alpine Italy (Br a m b i l l a 1970), South-alpine Switzerland (Wi e­d e n ­m a y e r 1977), Apennines (Sg u a zzo n i 1963), and Franco- ribs tend to become fastigate. The almost smooth venter nia (herein). does not show any indication of a keel. Within the umbili- Stratigraphic range: Late Pliensbachian (Stokesi to cus about six constrictions per whorl are discernible which Hawskerense subzones), possibly already present in the upper seem to disappear successively after a diameter of ca. Early Pliensbachian (Davoei Subzone), see Wi e d e n m a y e r (1977). In the Causses region this taxon is only reported from the Sub- 18 mm. nodosus to Gibbosus subzones (Me i s t e r 1989), but unknown yet The incompletely preserved second specimen (MAn- from the Spinatum Zone. 3115, Fig. 4) has a diameter of ca. 43 mm. It exhibits a Descriptions. – The first and smaller one of the two nearly completely preserved and uncrushed body chamber studied specimens (MAn-3111) measures 29.2 mm in di- (last half whorl). The phragmocone has been collapsed by ameter. The phragmocone of this completely preserved compaction processes and was lost along the first half of specimen is slightly compressed by compaction processes, the outer whorl, but is still visible within the open umbili- whereas its body chamber (last ½ whorl) does not show cus. The weak ribs run more or less radially over the flat- any diagenetic deformation (Fig. 3). However, on one of tened flanks of the body chamber and cross the rounded the flanks, the body chamber has lost a large angular piece venter along the last ¼ whorl in a fastigate style. Within of the shell affecting the entire flank and reaching from the open umbilicus, up to six constrictions per whorl are the peristome about 16 mm backwards. Obviously this discernible. The body chamber itself does not show any specimen was attacked by a predator, possibly a benthic constrictions. The morphology differs from the previously crustacean. Due to this large artificial opening of the body described specimens of Juraphyllites mimatensis espe- chamber, the measurements characterizing the shell mor- cially in the measurements of the umbilicus, which is sig- phology was taken along the smaller diameter of 24.2 mm nificantly narrower, with a relative width of about 16 % (see below). On the surface of the mould some relics of the versus 27–32 %, and the cross-section of the whorl which white aragonitic shell material are still preserved. The appears to be higher (ww/wh = 50 % versus 59–68 %). slightly compressed phragmocone shows the characteris- Only the umbilical elements of the last suture line are pre- tic juraphyllitid suture line (Fig. 3D) which matches per- served just at the beginning of the body chamber showing fectly with that of the lost holotype illustrated by d’Or b i g n y a characteristic juraphyllitic pattern formed by short am- (1865; Fig. 2) and the illustrations of the suture lines of the plitudes. From the suture line aspect, this specimen is neotype and additional material from the Causses provid- clearly distinguishable from the contemporaneous phyllo- ed by Jo l y (in Fi s c h e r 1994) and Jo l y (2000). Moreover, ceratid genus Zetoceras. In the latter, a similarly narrow also the measurements fall within the size ranges of the umbilicus is developed, but its suture line is much more specimens from the Causses (Jo l y 2000; see Tab. 1). The complex, with high amplitudes (see Jo l y 2000). The suture umbilicus of the specimen from Buttenheim is only slight- line, ribbing sculpture, and the lacking of a keel of this ly smaller, and the cross-section of the whorl is insignifi- specimen correspond with Juraphyllites mimatensis. How- cantly broader. Surprisingly, the holotype (Fig. 2) is the ever, due to the significantly smaller umbilicus and the closest one to our present finding although later authors higher whorl-section we prefer a cf.-determination of this stated that the illustration by d’Or b i g n y could be some- specimen. what generalized. 138 p a l a e o d i v e r s i t y 1, 2008

Fig. 4. Juraphyllites cf. mimatensis (d’Or b i g n y , 1845); Pliensbachian, Spinatum Zone, ? Apyrenum Subzone; Buttenheim, northern Bavaria, Germany; MAn-3115. – A. Lateral view. B. Ventral view. – Scale: 2 cm.

4. Palaebiogeography of Juraphyllites mimatensis finding of the keeled juraphyllitid Meneginiceras eximium (d’Or b i g n y , 1845) (Ha u e r , 1854) var. lariense (Me n e g h i n i , 1874) from the Lower Toarcian of the British Yorkshire coast. The inner- Without any doubt Juraphyllites mimatensis represents most whorls of Meneghiniceras have not developed the a Tethyan faunal element of the Late Pliensbachian which characteristic ventral keel of later ontogenetic stages and immigrated only during sea-level highstands and favour- hence they correspond morphologically well with the able currents into adjacent epicontinental basins, such as small and partly still unkeeled specimens of Ammonites the Causses area of Southern France and reached even tortisulcoides Qu e n s t e d t . As a consequence, Sc h w e i g e r t Southern Germany (see above). The records of this species (2005) interpreted the taxon Ammonites tortisulcoides in true Tethyan deposits are still remarkably scarce and Qu e n s t e d t , 1885 as a subjective junior synonym of Me- seem to be restricted to the Western Tethys. However, it is neghiniceras eximium (v. Ha u e r , 1854) due to the tenden- not unlikely that this species is much more widespread but cy to form a ventral keel with increasing diameters in most still unrecognized (see Wi e d e n m a y e r 1977: 37), because in of his specimens of Ammonites tortisulcoides under study. early ontogenetic stages this form is hardly distinguish- Interestingly, Mo n e s t i e r (1934) interpreted a specimen able form other juraphyllitids such as Meneghiniceras exi- from Swabia figured by Po m p e c kj (1893, pl. 1, figs. 10–11) mium (v. Ha u e r ) and Juraphyllites libertus (Ge m m e l l a ­ as belonging to d’Or b i g n y ’s species mimatensis, but he r o ). overlooked the keeled venter of this specimen. The pres- ence of a keel also in the lectotype of Ammonites tortisul- coides confirms the supposed synonymy of the two taxa. 5. Previously described juvenile Phylloceratina from Ke u p p & Sc ho b e r t (2008) recorded for the first time sev- the Upper Pliensbachian of Southern Germany eral very small, juvenile phylloceratid phragmocones, all of them less than 1 cm in diameter, from the Apyrenum Qu e n s t e d t (1885) described and illustrated tiny phyl- Subzone of Buttenheim. Following the argumentation loceratids from the Amaltheenton Formation (“Lias δ”) of given by Sc h w e i g e r t (2005), these innermost whorls were Gross-Eislingen (today: Eislingen) in Swabia as “Ammo- also determined as juveniles of the juraphyllitid Meneghin- nites tortisulcoides”. Other records assigned to this spe- iceras eximium (Ha u e r ) although a keel is not developed cies have been recorded from Southern Germany by Zw i e ­ in any of these tiny specimens. In Swabia, Meneghinic- s e l e (1898), En g e l (1908), Ur l i c hs (1977), Pl ü c k e b a u m eras eximium (v. Ha u e r ) occurs as an iteratively immigrat- (1985), and Sc h l e g e l m i l c h (1976). On the other hand, ing Tethyan guest in the Late Pliensbachian, but it was Ho w a r t h (1976) reported on a single excellently preserved never recorded from the Spinatum Zone. k e u p p & s c h w e i g e r t , juraphyllites m i m a t e n s i s i n so u t h e r n g e r m a n y 139

However, with the herein described specimens of Ju- Ho w a r t h , M. K. (1976): An occurrence of the Tethyan ammonite raphyllites mimatensis, a second juraphyllitid species is Meneghiniceras in the Upper Lias of the Yorkshire coast. – now reported which immigrated from the Tethyan into the Palaeontology, 19 (4): 73–777. Jo l y , B. (2000): Les Juraphyllitidae, Phylloceratidae, Neophyllo- epicontinental shallow sea of Southern Germany during ceratidae (Phyllocerataceae, Phylloceratina, Ammonoidea) the Late Pliensbachian. The juvenile stages of both taxa de France au Jurassique et au Crétacé. – Géobios, Mémoire appear to be indistinguishable. The same must be said spéciale, 23: 1–204. from the juveniles of another closely related juraphyllitid Ke u p p , H. (2005): Das Geheimnis der Spiralbänder: Farbmuster auf Ammonitengehäusen. – Fossilien, 22: 369–375. ammonite from the Tethyan Realm, Juraphyllites libertus Ke u p p , H. & Sc ho b e r t , J. (2008): Im Jugendstadium steckenge- (Ge m m e l l a r o ) (see list of synonyms in Fa n t i n i Se s t i n i blieben: Juvenile Meneghiniceraten aus Buttenheim. – Fos- 1974), which is rather evolute throughout ontogeny and silien, 25: 53–58. exhibits coarse ventrolateral ribs in the subadult and adult Ma c c h i o n i , F. (2001): Ammonites of the Domerian – Early Toar- cian in the Subbetic Zone and in the Umbria-Marche Apen- stages. nines. Taxonomy, taphonomy, biostratigraphy and paleobio- geography. 183 pp. PhD Thesis, Università di Perugia. Ma c c h i o n i , F. &. Me i s t e r , C. (2003): Ammonite biostratigraphy 6. Conclusions of some Mediterranean sections. 2: The succession of the Gola de F. Burano (Umbria-Machigiano Basin, Apennine), a reference section for Tethyan Domain. – Revue de Paléobi- In the Late Pliensbachian Tethyan juraphyllitids im- ologie, 22: 363–420. migrated in the Apyrenum and Hawskerense subzones of Ma i s c h , M. W. (2007): Dactylioceras (Eodactylites) holandrei Southern Germany and are recorded from Buttenheim in (d’Or b i g n y , 1844) (Ammonoidea, Dactylioceratidae) aus Franconia. These extremely weak immigration events dem Ober-Pliensbachium (Unterer Jura) von Bisingen (Baden-Württemberg). – Jahresberichte und Mitteilungen have not been recorded yet from Swabia, possibly due to des Oberrheinischen geologischen Vereins, Neue Folge, 89: collecting biases or more marly lithologies which are un- 49–58. favourable for ammonite preservation. Besides the jura- Me i s t e r , C. (1989): Les ammonites du Domérien des Causses phyllitids, represented by Juraphyllites (and cf.) mimaten- (France). – Cahiers de Paléontologie, 7: 7–80. Mo n e s t i e r , J. (1930): Observations nouvelles sur les ammonites sis and several undeterminable juveniles, no other Tethyan liasiques du Moyen Atlas Marocain. – Notes et Mémoires de immigrants are reported within the ammonite faunas of la Service géologique du Maroc, 8: 1–36. the Franconian Upper Pliensbachian absolutely predomi- Mo n e s t i e r , J. (1934): Ammonites du Domérien de la région Sud- nated by amaltheids (Pleuroceras ssp.). An unambiguous Est de l’Aveyron et de quelques régions de la Lozère à l’ex- clusion des Amalthéidés. – Mémoires de la Société géolo­ determination of juraphyllitids with diameters of less than gique de France, nouvelle série, 15: 7–70. 1 cm seems impossible. Op p e l , A. (1856–1858): Die Juraformation Englands, Frank­ reichs und des südwestlichen Deutschlands. – Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg, 12–14: 1–837. 7. References Or b i g n y , A. d e (1842–1851): Paléontologie Française. Descrip- tion zoologique et géologique de tous les animaux moll­ Br a m b i l l a , G. (1970): Le Ammoniti domeriana di Clivio (Vare- usques et rayonnés fossiles de France, comprenant leur ap- setto). – Atti della Società Italiana di Scienze Naturali e del plication à la reconnaissance des couches. Terrains ooli­ Museo Civico di Storia naturale in Milano, 110: 256–276. thiques ou jurassiques, 1, Céphalopodes. 624 pp.; Paris Du l l o , W.-C. (1981): Zur Erhaltung und Mikrostruktur der Am- (Masson). monitenschalen aus Unterstürmig (Ofr.). – Geologische Pa v i a , G. & Cr e s t a , S. (2002): Revision of Jurassic ammonites Blätter für Nordost-Bayern, 31: 75–83. of the Ge m m e l l a r o collections. – Quaderno Museo Geolo­ En g e l , T. (1908): Geognostischer Wegweiser durch Württem- gico “G. G. Gemmellaro”, 6: 406 pp. berg, 3rd edition. XXXI+645 pp.; Stuttgart (Schweizerbart). Pl ü c k e b a u m , M. (1985): Zur Stratigraphie und Paläontologie des Fa n t i n i Se s t i n i , N. (1974): Phylloceratina (Ammonoidea) del Oberen Pliensbach der Tongruben Marloffstein (Mfr.) und Pliensbachiano Italiana. – Rivista Italiana di Paleontologia e Unterstürmig (Ofr.) (unter besonderer Berücksichtigung der Stratigrafia, 80: 193–250. Ammonitenfamilie Amaltheidae Hyatt, 1867). 72 pp.; di- Fi s c h e r , J. C. (1994): Révision critique de la Paléontologie fran- ploma thesis, University of Erlangen [unpublished]. çaise de d’Or b i g n y . Vol. 1, Céphalopodes. 344 pp.; Paris Po m p e c kj , J. F. v. (1893): Beiträge zu einer Revision der Ammo- (Masson). niten des Schwäbischen Jura. – Jahreshefte des Vereins für Gu g e n b e r g e r , O. (1929): Paläontologisch-stratigraphische Stu- vaterländische Naturkunde in Württemberg, 49: 151–248. dien über den anatolischen Lias. – Neues Jahrbuch für Mi­ Qu e n s t e d t , F. A. (1885–1888): Die Ammoniten des Schwäbi­ neralogie, Geologie und Paläontologie, Beilage-Band, 62: schen Jura. 1140 pp.; Stuttgart (Schweizerbart). 235–300, 371–466. Ri c h t e r , A. E. (1977): Pyritisierte Ammoniten vom Fuße der Ha a r l ä n d e r , W. (1961): Erläuterungen zur Geologischen Karte Frankenalb. – Mineralien Magazin, 1 (1): 9–16. von Bayern 1 : 25000, Blatt Nr. 6434 Hersbruck. 76 pp.; Ri c h t e r , A. E. (1993): Die Tongrube Kalchreuth. – Fossilien, 10: München (Bayerisches Geologisches Landesamt). 18–24. Ho f f m a n n , R., Ke u p p , H. & Gr a d l , H. (2007): Zur Korrelation Ri c h t e r , A. E. (2000): Geoführer Frankenjura. 217 pp.; Augs- der Lias-Tongruben von Unterstürmig und Buttenheim burg (Ammon Rey Verlag) (Frankenalb). – Jahresberichte und Mitteilungen des Ober- Ri c h t e r , A. E. (2003): Richters Sammeltipp: Die Tongrube But- rheinischen geologischen Vereins, Neue Folge, 89: 37–48. tenheim. – Fossilien, 20: 95–103. 140 p a l a e o d i v e r s i t y 1, 2008

Sc h l e g e l m i l c h , R. (1976): Die Ammoniten des süddeutschen Ostracoda of the Upper Pliensbachian at the type locality Lias. 212 pp.; Stuttgart (G. Fischer). (Lias, SW-Germany). – Stuttgarter Beiträge zur Naturkunde, Sc h m i d t -Ka l e r , H. (2004): Wanderungen in die Erdgeschichte Serie B, 28: 1–13. (15): Das Walberla. 120 pp.; München (Pfeil). Vo i g t , E. (1968): Über Hiatus-Konkretionen (dargestellt an Sc h w e i g e r t , G. (2005): The occurrence oft he Tethyan ammonite Beispielen aus dem Lias). – Geologische Rundschau, 58: genus Meneghiniceras (Phylloceratina: Juraphyllitidae) in 281–296. the Upper Pliensbachian of SW Germany. – Stuttgarter Bei­ Wi e d e n m a y e r , F. (1977): Die Ammoniten des Besazio-Kalkes träge zur Naturkunde, Serie B, 356: 1–15. (Pliensbachian, Südtessin). – Schweizerische paläontologi­ Sg u a zzo n i , G. (1963): Fauna ammonitica die calcari selciferi sche Abhandlungen, 98: 1–131. metamorfici delle Alpi Apuane. – Palaeontographica Itali- Zw i e s e l e , H. (1898): Der Amaltheenton bei Reutlingen. 39 pp. ana, 57: 35–80. Inaugural-Dissertation, Universität Bern; Bern (Lack, Ur l i c hs , M. (1977): Stratigraphy, ammonite fauna and some Aesch­li­mann & Jost).

Addresses of the authors: Prof. Dr. He l m u t Ke u p p , Freie Universität Berlin, Department of Earth Sciences, Palaeontology, Malteserstraße 74-100, House D, 12249 Berlin, Germany E-mail: [email protected] Dr. Gü n t e r Sc h w e i g e r t , Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany E-mail: [email protected]

Manuscript received: 26.5.2008, accepted: 29.10.2008.