Early Jurassic (Upper Pliensbachian) gastropods from ...

Paläontologische Zeitschrift 2008, Vol. 82/1, p. 17–30, 31-03-2008

Early Jurassic (Upper Pliensbachian) gastropods from the Herforder Liasmulde (Bielefeld, Northwest Germany) SIEGFRIED SCHUBERT, Steinhagen; JOACHIM GRÜNDEL, Berlin & ALEXANDER NÜTZEL, München with 6 figures SCHUBERT, S.; GRÜNDEL, J. & NÜTZEL, A. 2007. Early Jurassic (Upper Pliensbachian) gastropods from the Herforder Liasmulde (Bielefeld, Northwest Germany). – Paläontologische Zeitschrift 82 (1): 17–30, 6 figs., Stuttgart, 31. 3. 2008. Abstract: An Early Jurassic (Upper Pliensbachian) gastropod fauna from the Herforder Liasmulde (Herford Lias

Syncline) near Bielefeld, Northwest Germany is described. This is one of the first detailed and illustrated reports of Pliensbachian gastropods from this area. Sixteen species are reported. Surface collecting yielded relatively large vetigastropods (pleurotomarioids amd trochomorphs). Bulk samples yielded abundant small caenogastropods (especially Levipleura blainvillei and Kalchreuthia frankei) and cylindrobullinid opisthobranchs (e.g. Cylindrobullina domeria). These small species are also abundant in the Pliensbachian of South Germany (N Bavaria, Franconia). Therefore, this characteristic Pliensbachian microgastropod assemblage was widespread in Germany and probably also in other comparable marine deposits of Central Europe (e.g. the Paris Basin). Keywords: Gastropoda • Jurassic • Pliensbachian • Amaltheenton Formation • Germany • taxonomy Kurzfassung: Eine unterjurassische (oberes Pliensbachium) Gastropodenfauna aus der Herforder Liasmulde nahe Bielefeld, Nordwestdeutschland, wird beschrieben. Es handelt sich um einen der ersten detaillierten und illustrierten Berichte von Gastropoden des Pliensbachiums aus dieser Region. Es werden 16 Arten belegt. Makroskopische Aufsammlungen lieferten vor allem großwüchsige Vetigastropoden (Pleurotomarioidea und Trochomorpha). Schlämmproben enthielten zahlreiche kleine Caenogastropoden (besonders Levipleura blainvillei und Kalchreuthia frankei) sowie cylindrobullinide Opisthobranchiata (z. B. Cylindrobullina domeria). Diese kleinwüchsigen Arten kommen auch im Pliensbachium Süddeutschlands (Franken) massenhaft vor. Dies zeigt, dass diese charakteristische Mikrogastropoden-Assoziation in Deutschland und vermutlich auch in anderen, vergleichbaren marinen Ablagerungen Zentral-Europas (z. B. im Pariser Becken) verbreitet war. eschweizerbartxxx

Schlüsselwörter: Gastropoda • Jura • Pliensbachium • Amaltheenton-Formation • Deutschland • Taxonomie

Introduction Upper Pliensbachian gastropods from NW Germany are not particularly well-studied and there are very few substantial studies (other than faunal lists) from the past decades. BRAUNS (1871) summarized and reported twelve species from the Amaltheenton Formation of N Germany but gave no illustrations. Recently SCHUBERT (2004a, b) listed a few gastropod species from the Herforder Liasmulde. But again, most of the material was not documented with illustrations (except Amberleya cf. subimbricata; SCHUBERT 2004a: pl. 7 fig. 3; KAEVER et al. 1976: 177, pl. 26). In contrast, Pliensbachian gastro-

pod faunas from Franconia (N Bavaria, S Germany) (NÜTZEL & KIEßLING 1997; GRÜNDEL & NÜTZEL 1998; NÜTZEL & HORNUNG 2002) and from NE Germany (GRÜNDEL 1999) were intensively studied during the last ten years. Thus, the study and documentation of the present gastropod fauna from the Herforder Liasmulde (Herford Lias Syncline) forms a considerable supplementation of our knowledge about faunal distribution in Central Europe. The goal of this paper is the documentation and illustration of this fauna as well as its taxonomic treatment. In addition, a detailed stratigraphic study including the occurrence of the gastropods is currently prepared by one of us (SCHUBERT).

Addresses of the authors: Siegfried Schubert, Magdeburger Straße 16, D-33803 Steinhagen. – Joachim Gründel, Institut für Geologische Wissenschaften, Fachbereich Paläontologie, Freie Universität Berlin, Malteser Str. 74–100, D-12249 Berlin; e-mail . – Alexander Nützel, Bayerische Staatssammlung für Paläontologie und Geologie, Richard Wagner Straße 10, D-80333 München; e-mail .

0031– 0220/08/0082–017

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© 2008 E. Schweizerbart’sche Verlagsbuchhandlung, D–70176 Stuttgart

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SIEGFRIED SCHUBERT et al.

Localities and preservation The material comes from the so-called Herforder Liasmulde (Herford Lias Syncline). The gastropods were collected by one of us (S. SCHUBERT) from various localities near Bielefeld (see SCHUBERT 2001, 2002, 2004a, b for descriptions and maps of the localities). The gastropods have a Late Pliensbachian (Domerian) age, including the zones of Amaltheus margaritatus and Pleuroceras spinatum (Amaltheus stokesi to Pleuroceras hawskerense subzones).

The preservation is quite variable. Some of the vetigastropods are well-preserved. Especially preservation in the concretions is good or very good. However, insufficient pyritization as well as steinkern preservation are also common. In the shales, the preservation is commonly poor. Moreover, the shales are difficult to disaggregate. However, one sample from a clay-pit near Jöllenbeck (Westerenger Straße I, NW exit of Jöllenbeck) yielded abundant microgastropods and some relatively well-preserved specimens. This sample is from the Amaltheus margaritatus Zone (Amaltheus gibbosus Subzone). The Amaltheenton Formation consists of thick, grey to blackish marine mudstones. They are wellknown for their rich ammonite faunas. These shales may also yield a rich benthic fauna which is, however, less conspicuous because it consists to a large part of smallsized species. The biota lived on (and partly in) a muddy soft bottom under fully marine conditions. Similar Pliensbachian shales from South Germany have been interpreted as being deposited under dysaerobic conditions with soft bottom conditions being one of the most important environmental constraints (NÜTZEL & KIEßLING 1997). Repository: All illustrated and some additional material is housed in the Naturkundemuseum Bielefeld (NKMB 32982–33017).

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Subclass Archaeogastropoda THIELE, 1925 Order Vetigastropoda SALVINI-PLAWÉN, 1980 Superfamily Pleurotomarioidea SWAINSON, 1840 Family Pleurotomariidae SWAINSON, 1840 Genus Pleurotomaria DEFRANCE, 1826 Pleurotomaria debuchii J.A. EUDESDESLONGCHAMPS, 1849 Figs. 2I–N

Fig. 1. – A: Location of Bielefeld in Germany, approximate extent of the Herforder Liasmulde shaded grey, see arrow. B: Extent of the Herforder Liasmulde, shaded grey, with locations of the collecting points. 1) Bardüttingdorf, middle margaritatus Zone to upper spinatum Zone; 2) Bielefeld-Jöllenbeck / Beukenhorst-I, lower margaritatus Zone to middle margaritatus Zone; 3) Bielefeld-Jöllenbeck / Beukenhorst-II, lower margaritatus Zone to middle margaritatus Zone; 4) Bielefeld-Jöllenbeck / Bodenheide, lower margaritatus-Zone to middle margaritatus Zone; 5) Bielefeld-Jöllenbeck / Dreeke, upper margaritatus Zone to lower spinatum Zone; 6) Bielefeld-Jöllenbeck / Westerenger Straße-I, upper margaritatus Zone; 7) Pödinghausen, Große Breede, lowermost margaritatus Zone.

* 1849 Pleurotomaria debuchii n. sp. J.A. EUDES-DESLONGCHAMPS: 90, pl. 16 fig. 1, pl. 17 fig. 5. 1997 Pleurotomaria (Talantodiscus) debuchii J.A. EUDESDESLONGCHAMPS, 1849 morphotype exertiuscula. – FISCHER & WEBER: 162, pl. 29 figs. 3, 4. See FISCHER & WEBER (1997) for more synonymy. Material: NKMB 33007, 33008.

Description: Shell low-spired trochoid; specimen illustrated in Figs. 2L–N 25 mm high and 41 mm wide;

Fig. 2. – A–H: Ptychomphalus expansus (SOWERBY, 1821). A–C. NKMB 33002; height 16 mm; width 28 mm. D–F. NKMB 33000; height 16.5 mm; width 28 mm. G–H. NKMB 33001; height 14 mm; width 19.5 mm. – I–N: Pleurotomaria debuchii J.A. EUDES-DESLONGCHAMPS, 1849. I–K. NKMB 33008; height 24 mm; width 38 mm. L–N. NKMB 33007; height 25 mm; width 41 mm.

Early Jurassic (Upper Pliensbachian) gastropods from the Herforder Liasmulde

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whorls asymmetrically convex with greatest width close to abapical suture; wide ramp present, demarcated by a rounded edge which is sometimes almost completely obscured; selenizone at about mid-whorl or slightly above on spire whorls, bordered by two spiral lirae, with distinct opisthocyrt lunulae; whorls evenly covered with numerous fine spiral threads; weak collabral axial ribs present on ramp, terminating at edge in small pustules; they become weaker during ontogeny and appear as strengthened growth lines on mature whorls; whorl face broadly rounded when joining weakly convex base; base with deep narrow umbilicus; growth lines prosocline on ramp, strongly curving backward at selenizone; growth lines below selenizone prosocyrt on whorl face and straight prosocline on base; aperture poorly preserved, wider than high, with rounded outer lip. Remarks: Our material closely resembles the Pliensbachian specimens from France as described and illustrated by FISCHER & WEBER (1997: pl. 29 figs. 3, 4) as morphotype exertiuscula EUDES-DESLONGCHAMPS. As far as we know, this or a similar species has not been reported from Germany. FISCHER & WEBER (1997) placed Pleurotomaria debuchii in the genus Talantodiscus FISCHER, 1885. However, Talantodiscus is a discoidal pleurotomariid and we are not convinced that species with a distinctly elevated spire should be placed in that genus. Family Eotomariidae WENZ, 1938 Genus Ptychomphalus AGASSIZ, 1837 (in J. SOWERBY 1812–1845)

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Ptychomphalus expansus (SOWERBY, 1821) Figs. 2A–H * 1821 Helicina expansa n. sp. J. SOWERBY: 129, pl. 273 figs. 1–3. 1997 Ptychomphalus expansus (J. SOWERBY). – FISCHER & WEBER: 160, pl. 26 figs. 1–3. 1998 Ptychomphalus expansus (J. SOWERBY). – GRÜNDEL & NÜTZEL: 66, pl. 1 figs. 6–9. See FISCHER & WEBER (1997) for more synonymy. Material: Three illustrated specimens: NKMB 33000, 33001, 33002.

Brief description: Rotelliform shell with slightly elevated spire, shallow ramp bordered by an adapical depression and a selenizone which forms the edge of the shell. Base with callus-filled umbilicus. Remarks: This characteristic shell is frequently mentioned in the literature. The original illustrations in SOWERBY (1821) closely resemble our material. SOWERBY’s (1821) material comes from the “Blue Lyas of Lyme in Dorsetshire”. However, study of the original material is needed to corroborate species identity because several Ptychomphalus species have been proposed which are relatively close to each other. Moreover, Ptychomphalus expansus seems to have a

wide intraspecific variabiliy and is relatively long ranging (Sinemurian to Pliensbachian according to FISCHER & WEBER 1997). Accordingly, the whorls are well separated from each other and may or may not have ledges. Even the spiral ornament varies from weak to strong and subsutural nodules may or may not be present. Several Ptychomphalus species from the Pliensbachian of S Germany have been described (we have material from Buttenheim and Kalchreuth at hand). A detailed description of material from Kalchreuth (S Germany, Franconia) was given by GRÜNDEL & NÜTZEL (1998). KUHN (1936) described four Ptychomphalus species from Franconia but did not report P. expansus. However, discrimination of these S German species is seemingly not well-founded and a reinvestigation of the original material is warranted. Superfamily Trochoidea RAFINESQUE, 1815 Family uncertain Genus Tylotrochus KOKEN, 1896 Tylotrochus subimbricatus (D’ORBIGNY, 1850) (n. comb.) Figs. 3A–G 1821 Trochus imbricatus n. sp. SOWERBY: 129, pl. 273 figs. 1–3. 1850 Trochus subimbricatus nom. nov. D’ORBIGNY: 267. 1871 Trochus imbricatus SOWERBY. – BRAUNS: 269. 1909 Trochus imbricatus SOWERBY. – BRÖSAMLEN: 209, pl. 17 figs. 19–21. 1976 Amberleya subimbricata (ORBIGNY). – KAEVER, OEKENTORP & SIEGFRIED: 177, pl. 26 fig. 6. 1990 Amberleya imbricata (SOWERBY 1854 [sic]) var. subimbricata (D’ORBIGNY). – HICKMAN & MCLEAN: 76, fig. 38B. 2004 Amberleya subimbricata (ORBIGNY). – SCHUBERT: 19, 23 (faunal lists). 2004 Amberleya cf. subimbricata (ORBIGNY). – SCHUBERT: pl. 7 fig. 3. See BRÖSAMLEN (1909) and BRAUNS (1871) for additional possible synonymy. Material: NKMB 33006 (Figs. 3D–G); height 22 mm, width 22.5 mm. Three additional illustrated specimens: NKMB 32993 (Fig. 3A), 32984 (Fig. 3B); 32982 (Fig. 3C). In addition several other specimens are present in the collection SCHUBERT.

Fig. 3. – A–G: Tylotrochus subimbricatus (D’ORBIGNY, 1850). A. NKMB 32993; height 21.5 mm; width 18 mm. B. NKMB 32984; height 36 mm; width 25 mm. C. NKMB 32982; height 22 mm; width 20 mm. D–G. NKMB 33006; height 22 mm; width 22.5 mm. – H–I: Eucycloscala dunkeri (GOLDFUSS, 1844), NKMB 33003; height 8 mm; width 8 mm. – J: Eucycloscala elegans (MÜNSTER, 1844), NKMB 32990; height 15 mm; width 12 mm. – K: Eucycloscala elegans (MÜNSTER, 1844), NKMB 32991; height 11 mm; width 9 mm. – L–M: Ooliticia? cyclostoma (BENZ, 1832), NKMB 33005; height 20 mm; width 15 mm.


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Description: Shell broad, conical; one of the illustrated specimens (Fig. 3B) is 36 mm high and 25 mm wide; apex and protoconch missing or corroded; flanks almost straight; whorls slightly convex, with pronounced keel just above lower suture which forms edge between whorl side and base; sutures very distinct due to this keel; whorl and base covered with numerous spiral ribs which bear or consist of abundant small nodules; 10–12 ribs present on whorl face; shell also covered with two systems (one prosocline, the other opisthocline) of linear non-collabral furrows which form a rhomboid scaly pattern (probably an artefact or trace fossil); base moderately convex; growth lines straight, prosocline only curving forward near the adapical suture; aperture simple, broad, oval, round. Remarks: Tylotrochus subimbricatus is a relatively often cited species (as “Amberleya” subimbricata ORBIGNY, 1858, replacement name for Trochus imbricatus SOWERBY, 1821). However, according to SOWERBY’S original illustration there seem to be some differences to our material. SOWERBY’s specimen is distinctly more slender and lacks nodular ornament. Moreover, the angulation of the whorls lies well above the suture in “Amberleya” subimbricata. However, A. KAIM was so kind to send us photos of English specimens from the Natural History Museum, London, of T. subimbricata (including the specimen illustrated by HICKMAN & MCLEAN (1990: 76, fig. 38B) and these specimens seem to be relatively close to our material. The specimen that was illustrated by HICKMAN & MCLEAN (1990: 76, fig. 38B) as “Amberleya imbricata (SOWERBY 1854 [sic]) var. subimbricata (D’ORBIGNY)” from the Lower Lias of England is very similar to our material and may be conspecific. However, this specimen from England also differs considerably from SOWERBY’S (1821) illustration of Trochus imbricatus. Even “Amberleya” subimbricata sensu QUENSTEDT (1858), BRÖSAMLEN (1909) and HÄGELE (1997) is more slender and in addition these species have fewer but well-defined spiral ribs. However, as in our material, they have small nodes on the spirals. Our material of Tylotrochus subimbricatus has a rhomboid scaly pattern on the teleoconch whorls. This rhomboidal structure is probably not a primary shell ornament but an artefact. J.A. TODD (London) (2006, written communication) suggests that it possibly represents attachment scars of hydrozoan colonies (see also TODD 1993; SZABO 1992). If it represented primary teleoconch ornament, it would be almost unique in trochomorph gastropods. The Permian species “Trochus” adrianensis GEMMELLARO, 1889 has a somewhat similar ornament (see GRECO 1937) but differs in shape and details of the ornament. The genus Tylotrochus KOKEN, 1896 is based on the Late Triassic type species Tylotrochus konincki (HÖRNES, 1856) which differs from Tylotrochus subimbricatus in having a distinct axial ornament in addition to the spiral ornament, so that a reticulate ornament is

formed (KOKEN 1896, 1897). Otherwise its shape and the presence of a suprasutural keel are shared with Tylotrochus subimbricatus. Even Tylotrochus keuppi BANDEL, GRÜNDEL & MAXWELL, 2000 from the Early Jurassic of New Zealand has a much more pronounced axial ornament. The early ontogenetic shell of Tylotrochus subimbricatus is unknown as is the early ontogeny of the type species of the type species of Tylotrochus. However, Tylotrochus keuppi lacks strong axial ribs on the early teleoconch and is therefore not closely related to the Eucyclidae. As far as we know, this characteristic species is absent in the Amaltheenton Formation of Franconia. Genus Eucycloscala COSSMANN, 1895 Eucycloscala dunkeri (GOLDFUSS, 1844) Figs. 3H–I * 1844 Turbo dunkeri n. sp. GOLDFUSS: 95, pl. 193 figs. 11a–b. ? 1936 Eucyclus dunkeri (MÜNSTER). – KUHN: 289, pl. 9 figs. 30–31. 1998 Eucyclus dunkeri (GOLDFUSS). – GRÜNDEL & NÜTZEL: 68, pl. 2 figs. 6–7. Material: One specimen, NKMB 33003.

Description: Shell trochiform, consisting of about five whorls, 8 mm high and wide; whorls moderately convex with ornament of three spiral rows of small but distinct pustules and numerous thread-like axial ribs.

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Remarks: The present specimen seems to agree well with material from Franconia (S Germany) where the type material of this species comes from (GOLDFUSS 1844; GRÜNDEL & NÜTZEL 1998). This is to our knowledge the first report of Eucycloscala dunkeri from the Herforder Liasmulde. Eucycloscala elegans (MÜNSTER, 1844) Figs. 3J–K * 1844 Turbo elegans MÜNSTER in GOLDFUSS: 94, pl. 193 figs. 10a–b. 1844 Turbo venustus MÜNSTER in GOLDFUSS: 94, pl. 193 figs. 9a–b. 1909 Eucyclus elegans (MÜNSTER). – BRÖSAMLEN: 256, pl. 20 figs. 7a–b. part 1936 Eucyclus elegans (MÜNSTER). – KUHN: 288, pl. 9 fig. 32. 1936 Eucyclus venustus (MÜNSTER). – KUHN: 288, pl. 9 figs. 19, 20, 22, 33. v 1997 Eucyclus elegans (MÜNSTER). – NÜTZEL & KIEßLING : 389, pl. 34 figs. 1–4. 1997 Eucyclus elegans (MÜNSTER). – HÄGELE: 63 (with illustration). v 1998 Eucyclus elegans (MÜNSTER). – GRÜNDEL & NÜTZEL: 389, pl. 2 figs. 4–5. Material: NKMB 32990, 32991.


Remarks: Two relatively high-spired turbiniform shells with convex whorls closely resemble the south German type material of Eucycloscala elegans. The whorls are ornamented with dominant spiral ribs. The earliest whorls seem to have a more knobby ornament. Ooliticia? cyclostoma resembles these specimens. However, Ooliticia? cyclostoma has much more spiral ribs (see below).

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Subclass Neritimorpha KOKEN, 1896 “Nerita” reticulata (MÜNSTER, 1844) Figs. 4D–F * 1844 Pileopsis reticulata n. sp. MÜNSTER in GOLDFUSS: 11, pl. 168 fig. 8. Material: A single specimen NKMB 33004.

Genus Ooliticia COSSMANN, 1894 Ooliticia? cyclostoma (BENZ, 1832) Figs. 3L–M, 4A–C * 1832 Turbo cyclostoma BENZ in ZIETEN: 45, pl. 33 figs. 4a–b. 1844 Turbo cyclostoma ZIETEN. – GOLDFUSS: 94, pl. 197 figs. 7a–b. 1852 Turbo cyclostoma ZIETEN. – QUENSTEDT: 420, pl. 33 fig. 35. 1854 Turbo cyclostoma ZIETEN. – OPPEL: 103, pl. 3 fig. 24. 1856 Turbo cyclostoma ZIETEN. – QUENSTEDT: 156, 195, pl. 19 figs. 27–28. 1876 Turbo cyclostoma BENZ. – TATE & BLAKE: 344, pl. 9 figs. 19–20. 1936 Turbo cyclostoma BENZ. – KUHN: 286, pl. 8 fig. 9. 1997 Ooliticia cyclostoma (BENZ, 1832). – HÄGELE: 68, text-fig. p. 68 lower left. Material: Four specimens: NKMB 32986, 32988, 32992, 33005.

Description: Medium-sized turbiniform shell, relatively high-spired; whorls distinctly convex; mature teleoconch whorls with numerous spiral and axial ribs; spiral ribs stronger than axials; intersections slightly nodular; axial ribs more and more reduced on mature teleoconch whorls. eschweizerbartxxx

Remarks: Ooliticia? cyclostoma is a widespread species in the margaritatus Zone (and spinatum Zone?) of Middle and Western Europe. Judging from the descriptions and illustrations in BENZ (in ZIETEN 1832), Turbo cyclostoma BENZ and Phasianella paludinaeformis SCHÜBLER (in ZIETEN 1832) represent synonyms (as was recognized previously by QUENSTEDT 1852). Phasianella paludinaeformis has page priority. However, since GOLDFUSS (1844) the name T. cyclostoma is used for this characteristic species throughout. Turbo cyclostomoides KOCH & DUNKER, 1837, described from North Germany, is probably a synonym. Likewise, Amberleya callipyge WILSON, 1887 and Littorina? chartroni COSSMANN, 1908 represent synonyms because illustrations and descriptions of these species show no difference that would justify specific differences. However, Turbo cyclostoma sensu BRÖSAMLEN (1909) is more slender, has more convex whorls and deeper sutures so that species identity is questionable. The assignment of Ooliticia? cyclostoma to the genus Ooliticia is questionable because the type species of Ooliticia differs considerably from that species (NÜTZEL & GRÜNDEL in prep.).

Description: We have at hand a relatively large specimen (26 mm wide) with rapidly increasing whorls and very wide aperture with almost straight to slightly curved inner lip. The spire is very low and only slightly protruding. Large parts of the last whorl with shell broken away exposing a steinkern with distinctly prosocyrt growth lines. Part of the shell is still preserved and shows an ornament of strong growth lines and fine spiral furrows. Remarks: The present specimen agrees well with the specimens illustrated by MÜNSTER (in GOLDFUSS 1844). Nerita alternans TATE & BLAKE, 1876 from the spinatum Zone of Yorkshire is also very similar and could be a synonym. MÜNSTER (in GOLDFUSS 1844) stated that the type specimen comes from the “Liaskalk bei Banz” (Lias limestone near Banz). At this location, various limestone units of Pliensbachian to Toarcian age are exposed. The assignment to “Nerita” is of course only preliminary due to the insufficient knowledge of this species. Even the assignment to the Neritimorpha is uncertain because certain vetigastropods (e.g. Stomatellidae) and Caenogastropoda have a similar shape. Protoconch morphology and shell structure are needed for a better substantiated placement. However, this relatively large shell is very characteristic and obviously rare though geographically widespread (North and South Germany, England). Unidentified neritimorph Figs. 4G–H Material: One specimen NKMB 32997.

Description: The present specimen is a low-spired trochoid steinkern with a high whorl expansion rate. The

Fig. 4. – A–C: Ooliticia? cyclostoma (BENZ, 1832). A. NKMB 32986; height 18 mm; width 15 mm. B. NKMB 32988; height 15 mm; width 12 mm. C. NKMB 32992; height 11.5 mm; width 10 mm. – D–F: “Nerita” reticulata (MÜNSTER, 1844), NKMB 33004, maximum width 26 mm. – G–H: Unidentified neritimorph, NKMB 32997, height 10 mm; width 13 mm. – I: Slab with Levipleura blainvillei and crinoids; width 48 mm; NKMB 32994. – J– O: Levipleura blainvillei (MÜNSTER, 1844). J. NKMB 32987–B, height 9 mm, steinkern with sculpture. K. NKMB 32998, height 11.5 mm, steinkern with sculpture. L. NKMB 32995–D, height 5.5 mm. M. From washed residue near Jöllenbeck, NKMB 33014; height 3.7 mm. N–O. From washed residue near Jöllenbeck, NKMB 33011; height 2.0 mm.

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Family Iravadiidae THIELE, 1928 Genus Kalchreuthia GRÜNDEL & NÜTZEL, 1998

whorls are round with deep sutures. The base is round with an evenly rounded transition to the whorl face. The aperture is broad oval. Remarks: This specimen has a lower whorls expansion rate than “Nerita” reticulata. Although it is too poorly preserved for any exact taxonomic assignment, this specimen documents the presence of an additional species in the present fauna. It could represent a neritimorph. Subclass Caenogastropoda COX, 1959 Family Zygopleuridae WENZ, 1938 Genus Levipleura GRÜNDEL & NÜTZEL, 1998 Levipleura blainvillei (MÜNSTER, 1844) Figs. 4I–O * 1844 Melania blainvillii n. sp. MÜNSTER in GOLDFUSS: 112, pl. 198 fig. 9. ? 1871 Cerithium blainvillii (MÜNSTER). – BRAUNS: 249. 1935 Zygopleura blainvillei (MÜNSTER, 1844). – KUHN: 483, pl. 18 fig. 7. 1995 Mesostrombus cf. vetustus (PHILLIPS, 1829). – SCHRÖDER: 37, pl. 6 figs. 18–20, pl. 15 fig. 3. v 1997 Zygopleura blainvillei (MÜNSTER, 1844). – NÜTZEL & KIEßLING : 390, pl. 34 figs. 5–7, pl. 35 figs. 1–2. v. 1998 Levipleura blainvillei (MÜNSTER 1844). – GRÜNDEL & NÜTZEL: 73, pl. 3 figs. 8–10. 1999 Katosira blainvillei. – HARRIES & LITTLE: 46, fig. 4. v 1999 Levipleura blainvillei (MÜNSTER 1844). – GRÜNDEL: 643, pl. 6 figs. 3–5. v 2002 Levipleura blainvillei (MÜNSTER 1844). – WEITSCHAT & GRÜNDEL: 39, 41, figs. 1–5, 11. v 2002 Levipleura blainvillei (MÜNSTER 1844). – NÜTZEL & HORNUNG: 56, pl. 1 fig. 1. eschweizerbartxxx

Material: Numerous specimens, NKMB 32994, 32987, 32995, 32998 from washed residue from Jöllenbeck: NKMB 33011, 33014, 33015.

Remarks: This species has a high-spired, small (about or less than 10 mm) shell with numerous axial ribs. The larval shell is multi-whorled and smooth. The present material from the Herforder Liasmulde agrees well with the material from South Germany (Franconia) as reported by MÜNSTER (in GOLDFUSS 1844), NÜTZEL & KIEßLING (1997) and GRÜNDEL & NÜTZEL (1998). BRAUNS (1871) mentioned that Levipleura blainvillei is present in the Harz foreland (N Germany). However, it has not been reported from the Herforder Liasmulde. MCDONALD & TRUEMAN (1921) reported Levipleura cf. blainvillei from the Liassic of England. It was also listed by HARRIES & LITTLE (1999) from the Pliensbachian of England. Levipleura blainvillei was found by surface collecting and is also very abundant in microsamples from Jöllenbeck (washed residues). Similarly, this species is one of the most abundant gastropods in South Germany (Franconia) (NÜTZEL & KIEßLING 1997; GRÜNDEL & NÜTZEL 1998).

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Kalchreuthia frankei (KUHN, 1936) Figs. 5A–B v* 1936 Pseudomelania frankei n. sp. KUHN: 294, pl. 13 fig. 11. v 1997 Tournoueria cf. frankei (KUHN). – NÜTZEL & KIEßLING : 393, pl. 35 figs. 5–7. v 1998 Kalchreuthia frankei (KUHN). – GRÜNDEL & NÜTZEL: 72, pl. 3 figs. 2–3. Material: Numerous specimens from a washed residue from Jöllenbeck; illustrated specimens NKMB 33012, 33013.

Remarks: This small caenogastropod was first reported from Franconia (South Germany) where it is locally very abundant (NÜTZEL & KIEßLING 1997; GRÜNDEL & NÜTZEL 1998) (detailed descriptions in these publications). Kalchreuthia frankei is characterized by a smooth and slender shell with a blunt protoconch. It is so small that it can usually only be found in microsamples. Kalchreuthia frankei is reported here from the Herforder Liasmulde for the first time. Family Aporrhaidae GRAY, 1850 Genus Dicroloma GABB, 1868 Dicroloma? sp. Fig. 5C Material: One specimen NKMB 32996.

Description: The juvenile specimen comprises about seven whorls, is 9 mm high and 4.2 mm wide; earliest whorls (probably larval shell) of about four to five convex whorls, somewhat greater apical angle than later whorls; early whorls without visible ornament; about fifth to sixth whorl with two faint spirals at about midwhorl bearing nodular ornament; last two whorls with distinct keel at mid-whorl, increasing in strength on last whorl; second keel forms edge at transition to base, covered by preceding whorl in earlier part of shell. Remarks: This juvenile specimen clearly represents an aporrhaid but it is too incomplete and poorly preserved for a safe generic or species identification. It resembles early ontogenetic shells of the Toarcian/Aalenian species Dicroloma? subpunctata (MÜNSTER 1844) (e.g. GRÜNDEL 1998). The present specimen represents one of the oldest members of the family Aporrhaidae in Germany. Family Carinariidae DE BLAINVILLE, 1818 Genus Tatediscus GRÜNDEL, 2001 Tatediscus aratus (TATE, 1870) Figs. 5F, G *

1870 Straparolus aratus n. sp. TATE: 404, pl. 26 fig. 21. 2001 Tatediscus aratus (TATE 1870). – GRÜNDEL: 60, pl. 5 figs. 5–8. See GRÜNDEL (2001) for more synonymy.

Material: One specimen NKMB 32985.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58

26 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58


Remarks: We have at hand a single well-preserved specimen from a concretion which agrees well with TATE’s (1870) illustration. It is a small planispiral shell with a strong ornament of broad, round axial ribs. Tatediscus aratus was reported from South Germany by BRÖSAMLEN (1909). This is to our knowledge the first report of this species from North Germany. Tatediscus aratus could represent a planktonic gastropod. Subclass Heterostropha FISCHER, 1885 Order Allogastropoda HASZPRUNAR, 1985 Superfamily Mathildoidea DALL, 1889 Family Gordenellidae GRÜNDEL, 2000 Genus Gordenella GRÜNDEL, 2000 Gordenella? sp. Fig. 5E Material: One specimen NKMB 32989.

Description: We have at hand a single poorly preserved specimen. It is high–spired, slender with numerous whorls and 22 mm high. The earliest preserved whorls are convex and have two spiral ribs on the lower part of the whorl. At least one addition weak spiral is present on the upper part of the whorl. Later whorls are only weakly convex to almost straight. The primary spiral ribs get closer to the abapical suture. Above them, at least three additional spiral ribs appear which are distinct but weaker than the primary spiral ribs. The growth lines are opisthocyrt. Intersections of spiral ribs and growth lines bear small pustules.

Order Opisthobranchia MILNE-EDWARDS, 1848 Familie Cylindrobullinidae WENZ, 1938 Genus Cylindrobullina AMMON, 1878 Cylindrobullina domeria (GRÜNDEL & NÜTZEL, 1998) Figs. 5H–I v part 1935 Actaeonina moorei DUM. – KUHN: 487, pl. 18 fig. 10 (non fig. 20). 1995 Actaeonina sp. – SCHRÖDER: 66, pl. 12 figs. 10– 13, pl. 15 fig. 17. v 1997 Actaeonina submoorei (KUHN). – NÜTZEL & KIEßLING : 395, pl. 36 figs. 1–7. v * 1998 Actaeonina domeria n. sp. GRÜNDEL & NÜTZEL 1998: 77, pl. 5 figs. 2–4. v 1999 Actaeonina domeria GRÜNDEL & NÜTZEL 1998. – GRÜNDEL: 656, pl. 9 figs. 6–9. v 2002 Cylindrobullina domeria (GRÜNDEL & NÜTZEL 1998). – NÜTZEL & HORNUNG: 56, pl. 1 fig. 3 Material: Numerous specimens from a washed residue from Jöllenbeck; illustrated specimen NKMB 33009.

Remarks: The genus was redefined by GRÜNDEL (2005) and placed in the Gordenellidae.

Remarks: Cylindrobullina domeria is a small-sized species with a medioaxial (terminology after SCHRÖDER 1995) larval shell (protoconch) and a broad subsutural ramp. Cylindrobullina domeria was described and discussed in detail by NÜTZEL & KIEßLING (1997) (as Actaeonina submoorei) and GRÜNDEL & NÜTZEL (1998). The holotype comes from the Sendelbach near Bamberg (North Bavaria, Franconia). It is one of the most abundant gastropods from the Amaltheenton Formation of Franconia. Some specimens from Jöllenbeck are well enough preserved for a substantiated species identification with the type material from South Germany (Franconia). Abundant steinkerns from the same microsamples from Jöllenbeck are probably conspecific. As in Franconia, Cylindrobullina domeria seems to be one of the most abundant species in the Herforder Liasmulde. This is the first report of Cylindrobullina domeria from this area.

Turritelloidea? sp.

Genus Ovactaeonina COSSMANN, 1895

eschweizerbartxxx

Remarks: The size and the high number of whorls and especially the ontogenetic change of the whorl profile and the ornament suggest that this species represent the genus Gordenella.The preservation is too poor for a species identification. Genus Turritelloidea WALTHER, 1951

Fig. 5D Material: One specimen NKMB 32999.

Description: The only specimen is a teleoconch fragment of about eight whorls that is 17 mm high. It is highspired and slender. The whorl face is evenly convex with distinct sutures. The whorls are entirely covered with several (up to 10) fine but distinct spiral ribs. The base is weakly convex. Remarks: This specimen resembles other Turritelloida species in size, shape and ornament of the mature ontogenetic stages. However, in the absence of diagnostic early ontogenetic characters, a species identification as well as a certain generic assignment are impossible.

Ovactaeonina sp. Figs. 5J–L Material: Several specimens from a washed residue from Jöllenbeck, illustrated specimen NKMB 33010.

Remarks: We illustrate a small heterostrophic opisthobranch from Jöllenbeck which lacks a distinct subsutural ramp that would be typical for Cylindrobullina. Such shells have been repeatedly reported from the Amaltheenton Formation of Germany (e.g. SCHRÖDER 1995; NÜTZEL & KIEßLING 1997; GRÜNDEL & NÜTZEL 1998). The preservation of the present specimens is too poor for species identification.


27

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Fig. 5. – A–B: Kalchreuthia frankei (KUHN, 1936), from washed residue near Jöllenbeck. A. NKMB 33013, height 1.2 mm. B: NKMB 33012, height 1.2 mm. – C: Dicroloma? sp., NKMB 32996, height 9 mm. – D: Turritelloidea? sp., NKMB 32999, 17 mm. – E: Gordenella? sp., NKMB 32989, height 22 mm. – F–G: Tatediscus aratus (TATE, 1870), NKMB 32985. – H–I: Cylindrobullina domeria (GRÜNDEL & NÜTZEL, 1998), from washed residue near Jöllenbeck, NKMB 33009, height 1.2 mm. – J–L: Ovactaeonina sp., from washed residue near Jöllenbeck, NKMB 33010. J. height 1.0 mm; K. height 1.0 mm. L. height 0.6 mm.

Driftwood with possible gastropod spawn Fig. 6

A piece of driftwood (7 cm wide, 12 cm long, NKMB 32984) from the Upper gibbosus Subzone of Dreeke has numerous small pyritic hemispheres attached. The hemispheres are flat and have a diameter of 0.35 mm to somewhat more than 1 mm. The hemispheres are commonly deformed with sunken structures. Similar Early Jurassic spawn-like structures on a bivalve shell have been reported by KAISER & VOIGT (1977). A very similar driftwood with attached pyritic hemispheres from the Bielefeld area (same age) was reported and discussed by RIEGRAF & SCHUBERT (1991). We refer to the discussion of these structures in these publications. The deformation of the hemispheres

shows that these objects were still flexible before pyritization. This supports the view that they represent invertebrate (possibly gastropod) spawn and that pyritization was very rapid.

Discussion of the gastropod fauna This paper represents the first encompassing treatment of gastropods from the Upper Pliensbachian Amaltheenton Formation of NW Germany in which specimens are illustrated and which is therefore easier falsifiable than previous faunal lists from that region. This collection certainly overrepresents vetigastropods because the present species are relatively large and therefore preferably picked up by surface collecting. Some of these

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Fig. 6. Wood with attached pyritized possible gastropod spawn; NKMB 32983. – A–C: details from D, each 9 mm wide. – D: Piece of wood 7 cm wide, 12 cm long. eschweizerbartxxx

large vetigastropods (Pleurotomaria debuchii, Tylotrochus subimbricatus) have not been reported from the rich gastropod faunas of Franconia (South Germany). In contrast, Ptychomphalus expansus seems to be widespread and occurs in Franconia as well as in England. The gastropod fauna of the here examined micro-samples (washed residues) is dominated by small caenogastropods (Levipleura blainvillei and Kalchreuthia frankei) and by several small opisthobranch species (especially Cylindrobullina domeria). This microgastropod assemblage is definitely similar to those of Franconia (South Germany) (NÜTZEL & KIEßLING 1997; GRÜNDEL & NÜTZEL 1998; NÜTZEL & HORNUNG 2002) and some of these species are also present and abundant in other parts of North Germany (GRÜNDEL 1999; WEITSCHAT & GRÜNDEL 2002). This characteristic assemblage of small gastropods has been largely unrecognized until recently. Its presence in the Herford Lias Syncline is an important step towards a better understanding of faunal distribution of the Lias of Central Europe. As in South Germany, gastropods dominate the benthic fauna in terms of species richness and abundance. Further considerations are somewhat hampered by insufficient knowledge (lack of modern taxonomic

work) of Jurassic microgastropods from England and France. The knowledge of Pliensbachian faunas from Europe is also an important step towards a better understanding of the end-Pliensbachian mass extinction event and associated onset of anoxia.

Acknowledgements We thank Andrzeij Kaim (Tokyo and Warsaw) and Sven Nielsen (Kiel) for their careful reviews and Mrs Neufert (Erlangen) for technical assistance.

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Manuskripteingang / manuscript received 19. 12. 2006; Manuskriptannahme / manuscript accepted 10. 10. 2007.